Hordeum chilense resistance to powdery mildew and its potential use in cereal breeding

Summary Hordeum chilense is a wild barley with high crossability with Triticum, Hordeum and Secale. Its amphiploid with wheat, tritordeum, has potential as a new crop. H. chilense is highly resistant to the powdery mildew diseases of both wheat and barley. Whereas tritordeum is resistant to barley powdery mildew, its reaction to wheat powdery mildew is similar to that of its wheat parent. However H. chilense contributes to a reduced density of mildew colonies. This quantitative resistance of tritordeum is diluted at higher ploidy levels.     

The reaction of x Tritordeum and its Triticum spp. and Hordeum chilense parents to rust diseases

Summary Hexaploid and octoploid tritordeums and their parents Hordeum chilense and Triticum spp. were screened for resistance to isolates of wheat and barley yellow and brown rusts. All H. chilense lines were highly resistant to both wheat and barley brown rust, few lines were susceptible to wheat yellow rust while susceptibility to barley yellow rust was common. In general the resistance of tritordeum is predominantly contributed by the wheat parent and apparently the genes for resistance in H. chilense are inhibited in their expression by the presence of the wheat genome.Abbreviations WYR wheat yellow rust - WBR wheat brown rust - BYR barley yellow rust - BBR barley brown rust     

The contribution of Hordeum chilense to partial resistance of tritordeum to wheat brown rust

Summary Hexaploid and octoploid tritordeum and their Triticum spp. parents were studied in the seedling stage to compare their components of partial resistance to Puccinia recondita f.sp. tritici. The components studied were infection frequency, latency period and size of uredia. The non-host Hordeum chilense parent does not confer any increase of partial resistance to wheat brown rust to its amphiploids with wheat.     

Chromosomal location of resistance to Septoria tritici in Hordeum chilense determined by the study of chromosomal addition and substitution lines in `Chinese Spring' wheat

Hordeum chilense exhibits resistance to Septoria tritici. Addition and substitution lines of H. chilensein wheat were utilized in growth chamber and field experiments to determine which H. chilense chromosomes carry resistance genes. Resistance is conferred by gene(s) on chromosome 4 and, to a minor extent, by genes on chromosomes 5, 6 and 7. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genome discrimination and chromosome pairing in the Hordeum chilense × Aegilops tauschii amphiploid

Tritordeum (X Tritordeum Ascherson et Graebner) is a synthetic amphiploid belonging to the Triticeae tribe, which resulted from crosses between Hordeum chilense and wheat. It presents useful agronomic traits that could be transferred to wheat, widening its genetic basis. In situ hybridisation with total genomic DNA from H. chilense and cloned, repetitive DNA sequences (pTa71 and pAs1) probes were used to discriminate the parental origin of all chromosomes, to analyse the chromosome pairing and to identify the chromosomes in pollen mother cells (PMCs) at metaphase I of the tritordeum line HT251 (H^chH^chDD, 2n = 4x = 28). The H. chilense total genomic DNA and the ribosomal sequence pTa71 probes, allowed the unequivocal discrimination of the 14 chromosomes of H^ch genome-origin and the 14 chromosomes of D genome-origin. Chromosome pairing analysis revealed meiotic irregularities such as reduced percentage of PMCs with complete homologous pairing, high frequency of univalents, most of H. chilense-origin and a reduced frequency of intragenomic multivalents from both genomes. The H. chilense genome revealed high meiotic instability. After individual chromosome identification at metaphase I with the pAs1 probe, we found the occurrence of pairing between chromosomes of different homoeology groups. The possible interest of the tetraploid tritordeum in the improvement of other Triticeae species is also discussed.     

Genetic variation for carotenoid pigment content in the amphiploid Hordeum chilense × Triticum turgidum conv. durum

Hexaploid tritordeum, the amphiploid Hordeum chilense x Triticum turgidum conv. durum has a higher grain carotene content than durum wheat. In order to decide strategies for introgressing this character into durum wheat, the effect on the carotene content of tritordeum synthesized with H. chilense and durum wheat differing in carotene content was analysed. Carotene content was evaluated in 35 primary tritordeum lines and their parents, 27 H. chilense accessions and 19 durum wheat cultivars. Some amphiploids have either one barley or wheat parent in common. In general, the influence of H. chilense is more important than that of wheat in the amphiploid carotene content. Nevertheless, the interactions between both parents on the amphiploid carotene content are also important.     

Resistance to Septoria tritici in Hordeum chilense×Triticum spp. Amphiploids

The reaction of tritordeum and its Hordeum chilense and Triticum spp. parents to Septoria tritici was studied in field and seedling experiments. All H. chilense lines were highly resistant to all the isolates and did not allow pycnidia development. The ‘durum wheat isolate’ developed pycnidia only on durum wheats. The ‘breed wheat isolate’ was very virulent on bread wheat but also on the wild tetra-ploid wheats. The other two isolates were compatible with durum and bread wheat. All hexaploid tritordeums were highly resistant both in the field and the seedling experiments. Some octoploid tritordeums allowed pycnidial development, but at much lower levels than their wheat parent. Resistance in tritordeum was not associated with plant stature and only in octoploid tritordeum was association of resistance with late maturity detected.     

Chromosomal localization of genes for carotenoid pigments using addition lines of Hordeum chilense in wheat

×Tritordeum sp. (Ascherson et Graebner) is the amphiploid obtained after chromosome doubling of hybrids between Hordeum chilense (Roem. et Schult.) and diploid, tetraploid or hexaploid wheats. Tritordeums have consistently higher carotenoid pigment contents than durum or bread wheat. Two distinct H. chilense accessions (used for the synthesis of tritordeum) were analysed for this trait. The chromosomal localization of the genes coding the ability of H. chilense to increase the carotene content of wheat were carried out using two sets of wheat- H. chilense addition lines. The a arm of chromosome 7H^ch is proposed to be responsible for the high carotene content in tritordeum. The implication of this finding in wheat breeding is discussed.     

Short Communication Resistance to common bunt in Hordeum chilense X Triticum spp. Amphiploids

The reaction of tritordeum and its Hordeum chilense and Triticum spp. parents to common bunt incited by Tilletia tritici were determined in field experiments. H. chilense accessions were very resistant, and durum wheats exhibited high to moderate levels of resistance. Conversely, bread wheats were highly susceptible. Resistance from H. chilense was expressed in the amphiploids, although the level of resistance was partially diluted at higher ploidy levels. Hexaploid tritordeums were immune to the disease; some infection was observed among the octo-ploids but at much lower levels than in their respective wheat parents.     

A physical map of chromosome 4Hch from H. chilense containing SSR, STS and EST-SSR molecular markers

The construction of a physical map of chromosome 4H^ch from Hordeum chilense containing molecular markers capable of detecting segments of this chromosome in a wheat background would be very useful for marker-assisted introgression of 4H^ch chromatin into both durum and common wheat. With this aim, the applicability of 106 barley chromosome 4H primers (62 SSRs and 44 STSs) to amplify markers showing polymorphism between H. chilense and both common or bread and durum wheat was investigated. Twenty-five SSR (40.3%) and six STS (13.6%) barley primer pairs consistently amplified H. chilense products. Eight SSR (12.9%) and four STS (9.1%) barley primers were polymorphic between H. chilense and both common and durum wheat, 10 of them (6 SSRs and 4 STSs) were located on chromosome 4H^ch using both the addition line of chromosome 4H^ch in Chinese Spring wheat and a tritordeum line (an amphiploid between H. chilense and T. turgidum) nullisomic for chromosome 4H^ch. Additionally, 18 EST-SSR barley markers previously located on chromosome 4H^ch were screened for polymorphism; 15 were polymorphic between H. chilense and both durum and common wheat. For physical mapping we used a ditelosomic tritordeum line for the short arm of chromosome 4H^ch and a tritordeum line homozygous for a 70% terminal deletion of the long arm of 4H^ch. A total of 25 markers (6 SSRs, 4 STSs and 15 EST-SSRs) were mapped to chromosome 4H^ch. Eight markers were allocated on the 4H^chS, eight were mapped in the 30% proximal region of 4H^chL and nine were on the 70% distal region of 4H^chL, respectively. Arm location on barley chromosome 4H was also carried out using both 4HS and 4HL ditelosomic addition lines in wheat. All markers mapped may have a role in marker-assisted introgression of chromatin segments of chromosome 4H^ch in both durum and common wheat backgrounds. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Development and cytogenetic characterisation of a double goat grass-barley chromosome substitution in tritordeum

×Tritordeum (Ascherson et Graebner, an amphiploid between Triticum turgidum conv. durum and Hordeum chilense), and chromosome substitution lines of tritordeum where chromosomes 2 H ^ch or 3 H ^ch H. chilense were replaced with chromosome 2 D of T. aestivum or 3 H ^v chromosome of H. vulgare, respectively, were used to assess the effect of specific chromosomes on the rachis. ×Tritordeum has brittle rachis while the 2 D(2 H ^ch) and 3 H ^v (3 H ^ch) substitution lines have non-brittle rachis. Both lines also have compact spikes, a character highly desirable for the improvement of tritordeum threshability. Different combinations of 2 D and 3 H ^v translocations were developed in tritordeum. In this article we present information on the identification and characterisation of all these introgression lines by the fluorescent in situ hybridisation.     

Somatic Embryogenesis and Plant Regeneration from Tritordeum

Regeneration of plants by somatic embryogenesis from immature embryos of hexaploid tritordeum (AABBH^chH^ch, amphiploid Hordeum chilense×Triticum turgidum conv. durum) and durum wheat (Triticum tergidum) was induced on MS medium supplemented with different 2.4-D concentrations. Well-defined embryoids were formed with a high frequency on the scutellar callus from 1 or 2 weeks onwards and plantlets were developed from them. In the best cases from one single explant more than 100 plants could be obtained. Plants were also regenerated by somatic embryogenesis from inflorescences of Hordeum chilense×Triticum turgiditm conv. durum hybrid and its respective hexa-amphiploid. With regard to callus induction and regenerative ability, evident differences between hexa- and octoploid (H. chilense×T. aestivum) tritordeum were found, the latter showing a very low response.     

Genealogical Identification of Hexaploid Tritordeum by Electrophoretic Separation of Endosperm Storage Proteins

Hexaploid Tritordeum (×Tritordeum Ascherson et Graebner) has been analyzed by SDS-PAGE, to test the efficiency of the method for confirming the parents of different lines, and for unequivocally discriminating between the lines. The results clearly show that endosperm storage proteins from both parents (durum wheat ——Triticum turgidum conv. durum Desf. em. M.K. —— and Hordeum chilense Roem. et Schulz.) are expressed in the endosperm of the amphiploid between them. Although the spectra of proteins from the parental species overlap on the gel, there are sufficient number of bands with distinct electrophoretic mobilities to identify unambiguously their parental origin. Furthermore, in the six lines of tritordeum analyzed, obtained from crosses between three accessions of H. chilense and four cultivars or lines of wheat, there was sufficient polymorphism amongst the parents, particularly amongst the H. chilense accessions, to enable both aims of the project to be met, that is, to confirm the genealogy of each tritordeum, and to discriminate between different lines. This study provides the basis of future investigation on the relationship between bread-making quality in tritordeum and the allelic composition of storage proteins.     

Chromosome and chromosomal arm locations of genes for resistance to Septoria glume blotch in wheat cultivar Cotipora

Summary Septoria glume blotch, caused by Stagonospora nodorum, is an important disease of wheat (Triticum aestivum). Separate genetic mechanisms were found to control flag leaf and spike resistance. Genes for resistance to S. nodorum were located on different chromosomes in the few wheat cultivars studied. These studies only partially agree on the chromosome locations of gene in wheat for resistance to S. nodorum, and chromosomal arm locations of such genes are not known. The objectives of this study were to determine the chromosome and chromosomal arm locations of genes that significantly influence resistance to S. nodorum in wheat cultivar Cotipora. Monosomic analysis showed that flag leaf resistance was controlled by genes on chromosomes 3A, 4A, and 3B whereas the spike resistance was controlled by genes on chromosomes 3A, 4A, 7A, and 3B (P=0.01). Additionally, genes on chromosomes 6B and 5A influenced the susceptibility of the flag leaf and spike reactions, respectively (P=0.01). Telocentric analysis showed that genes on both arms of chromosome 3A, and the long arms of chromosomes 4A and 3B were involved in the flag leaf resistance whereas genes on both arms of chromosome 4A, the short arm of chromosome 3A, and the long arm of chromosome 3B conferred spike resistance.     

Avoidance of leaf rust fungi in wild relatives of cultivated cereals

Summary Avoidance of rust fungi that was based on poor appressorium induction was previously found in Hordeum chilense. In the present study 95 accessions of Triticeae were screened for avoidance of Puccinia hordei. The percentage of appressorium formation per germinated spore ranged from 6 to 90%. On none of the 41 accessions of Aegilops, Agropyron, Elymus, Secale, Thinopyrum or Triticum studied was the rate of appressorium formation lower than 25%. Lower rates of appressorium formation were, however, found on accessions of wild barley species Hordeum brachyantherum, H. marinum, H. parodii and H. secalinum. Its implications in cereal breeding are discussed.     

Analysis of D-prolamins synthesized by the Hordeum chilense genome and their effects on gluten strength in hexaploid tritordeum

Hexaploid tritordeum is the amphiploid derived from the cross between Hordeum chilense and durum wheat. The storage proteins synthesized in the H^ch genome influence the gluten strength of this amphiploid. The D‐prolamins of H. chilense have been analysed by sodium dodecyl sulphate‐polyacrylamide gel electrophoresis with and without urea. A new locus named Glu‐H^ch3 has been detected. The effects of allelic variation at this locus on gluten strength, as measured the sodium dodecyl sulphate sedimentation test, were determined using seeds of 92 lines from a cross of two hexaploid tritordeum lines. Two allelic variants have been detected for this locus, which have shown different effects on gluten strength.     

Response of wheat cultivars to infection by Stagonospora nodorum isolates/mixture on detached and intact seedling leaves

Stagonospora nodorum blotch (SNB) of wheat caused by Stagonospora nodorum (Berk.) Castellani & E.G. Germano is among the major foliar diseases of wheat worldwide. The response of four wheat cultivars for reaction to infection by isolates/mixture of S. nodorum was tested on detached and intact seedling leaves in the laboratory and glasshouse experiments, respectively. The four wheat cultivars tested significantly differed from each other in response to infection by the isolates/mixture of S. nodorum. Similarly, the aggressiveness of the four isolates/mixture of the pathogen on the four wheat cultivars was significantly different as demonstrated by the parameters tested in the two experiments. However, no significant (p ≤ 0.05) isolate x cultivar interaction was observed for all the parameters tested in the two experiments. Highly significant positive or negative correlations were observed between parameters, i.e., disease severity (DS), incubation period(IP), and size of necrotic lesions (SNL), tested on detached and intact seedling leaves, respectively. This suggests that use of the detached leaf technique has considerable promise for quick screening of wheat cultivars against SNB. This revised version was published online in August 2006 with corrections to the Cover Date.     

Bread-making quality in hexaploid tritordeum with substitutions involving chromosome 1D

Hexaploid tritordeum, the amphiploid Hordeum chilense×Triticum turgidum, has potential for bread making. In order to estimate the potential of bread wheat chromosome 1D for improving the bread‐making quality of tritordeum, and the processing properties and agronomic performance of euploid tritordeum, (1H^ch)1D and (1A)1D substitution lines have been evaluated in field trials. No significant differences for agronomical traits were observed between the two substitution lines and the sister euploid tritordeum, except for the kernel weight of the (1H^ch)1D tritordeum substitution, which was lower than that of euploid tritordeum. Gluten strength, estimated by alveograph deformation energy (W), and loaf volume were substantially higher in both substitution lines than in the euploid tritordeum.     

Resistance against greenbug, Schizuphis graminum Rond., and Russian wheat aphid, Diuraphis noxia Mordvilko, in tritordeum amphiploids

A collection of tritordeum amphiploids (Hordeum chilense × Triticum turgadum) and their wheat parents were screened for resistance against the two main aphid pesis of cereals, the greenhug. Schizaphis graminum Rond. and ihe Russian wheat aphid (RWA) Diuraphis naxia Mord-vilko. Antixenosis. antibiosis and tolerance were evaluated in controlled environmental conditions using a. clone of greenbug biotypc C and a clone of RWA collected on pasta wheat. Tritordeum amphiploids pos-sess genetic resistance against greenbug and RWA; some of the lines tested were more resistant than the parental wheat line. Four principal components explained the resistance against both aphid species. The antixenosis shown against both pests was mainly contributed by their wheat parents. The antibiosis againsl both aphid species was obviously dependent on diflerent plant traits. The highest levels of antibiosis against the two aphids occurred in different amphiploids. Different genes are involved in the antibiotic reaction against the two aphids. The Tritordeum resistance to RWA is based on anlixenosis and ant-biosis since the tolerance trails were not independent of the other types of resistance. The level of tolerance shown to the greenbug was variable and appears to be controlled by differeni mechanisms. The tolerance to aphids shown by H. chilense is expressed in the amphiploids. but with some genomic interaction. Genes conferring resistance to aphids in H. chilensee could be incorporated into new cultivars of wheat to broaden their genetic base of resistance against greenbug and RWA.     

Effects of Hordeum chilense and Triticum Cytoplasms on Agronomical Traits in Hexaploid Tritordeum

The genome of Tritordeum, AABBH^chH^ch, was substituted into the cytoplasms of Triticum aestivum, T. turgidum and Hordeum chilense by repeated back-crossing to produce alloplasmic lines. This substitution did not greatly affect the characters studied, except yield per plot and fertile ears per plant, which were lower on T. turgidum cytoplasm. Cytoplasm from either H. chilense or T. aestivum could be used for breeding tritordeum.