Selection of U and M genome-specific wheat SSR markers using wheat–<Emphasis Type="Italic">Aegilops biuncialis</Emphasis> and wheat–<Emphasis Type="Italic">Ae. geniculata</Emphasis> addition lines

We selected wheat SSR markers specific to the U and M genomes of Aegilops species. A total of 108 wheat SSR markers were successfully tested on Ae. biuncialis (2n = 4x = 28, U^bU^bM^bM^b), on five wheat–Ae. biuncialis addition lines (2M^b, 3M^b, 7M^b, 3U^b and 5U^b) and on a wheat–Ae. geniculata (1U^g, 2U^g, 3U^g, 4U^g, 5U^g, 7U^g, 1M^g, 2M^g, 4M^g, 5M^g, 6M^g and 7M^g) addition series. Among the markers, 86 (79.6%) were amplified in the Ae. biuncialis genome. Compared with wheat, polymorphic bands of various lengths were detected on Ae. biuncialis for 35 (32.4%) of the wheat microsatellite markers. Three of these (8.6%) exhibited specific PCR products on wheat–Ae. biuncialis or wheat–Ae. geniculata addition lines. The primers GWM44 and GDM61 gave specific PCR products on the 2M^b and 3M^b wheat–Ae. biuncialis addition lines, but not on the 2M^g addition line of Ae. geniculata. A specific band was observed on the 7U^g wheat–Ae. geniculata addition line using the BARC184 primer. These three markers specific to the U and M genomes are helpful for the identification of 2M^b, 3M^b and 7U^g chromosome introgressions into wheat.     

Growth dependency of wild,primitive and modern cultivated wheat lines on vesicular-arbuscular mycorrhiza fungi

Summary Mycorrhizal colonization and growth dependency were studied at a single rate of phosphorous application in wild and cultivated primitive and modern wheats, inoculated with Glomus intraradices Schenck & Smith. Mycorrhizal colonization found in Triticum timopheevii var. araraticum (AAGG) was higher than that found in the other tetraploid wheats (AABB). Mycorrhizal dependency was higher in representatives of the D genome donor — Aegilops squarrosa, compared with representatives of the A and possible B genome donors T. monococcum and Ae. sharonensis, Ae. longissima and Ae. speltoides, respectively. The nature of response to VAM in hexaploid wheat was controlled by factors of the A and B genomes which are epistatic over those located in the D genome. The high mycorrhizal colonization and dependency which was found in T. timopheevii var. araraticum may indicate special genomic affinity possessed by the G genome of wheat in VAM interaction. Based on the 27 wheat lines and species tested in this study only low correlation between G. intraradices colonization and its contribution to plant growth can be suggested.     

Evaluation of a collection of wild wheat relative Aegilops geniculata Roth and identification of potential sources for useful traits

A collection comprising 157 Aegilops geniculata accessionsoriginating from different ecogeographical regions was established atENSA-INRA, Montpellier. The accessions were studied for physiologicaltraits related to drought and heat stress and screened for resistance tobarley yellow dwarf virus (BYDV) and rusts. Some accessions were alsotested for resistance to Hessian fly and cereal cyst nematodes (CCN). The study allowed to distinguish different adaptive strategies to theclimatic constraints encountered by Aegilops populations in theirregion of origin. They led to significant differences in biomass and grainproduction and should be taken into account in the utilisation of Ae.geniculata germplasm in wheat breeding programs. Two accessions withresistance to BYDV were found. Both originated from South of France. Theinterest of Ae. geniculata as a source of rust resistance was confirmedand accessions with resistance to the three rusts were identified. Highresistance against populations of Heterodera avenae and H.latipons was found in accessions from Spain, Bulgaria, Jordan and Tunisia.Sources of resistance to Hessian fly were also identified.     

The effect of transfers of alien genes for leaf rust resistance on the agronomic and quality characteristics of wheat

Summary Nine transfers of leaf rust (Puccinia recondita Rob. ex Desm.) resistance to wheat (Triticum aestivum L.) from Agropyron elongatum Host. Beauv., Triticum speltoides Tausch and rye (Secale cereale L.) were backcrossed up to 10 times to commercial wheat cultivars. The objective was to study the effect of the transfers on agronomic and quality characters and to make them available in desirable genetic backgrounds. The results varied greatly for different transfers. In four cases no promising material was obtained even after nine backcrosses. However, for the remaining five transfers material with potential as a new cultivar was obtained.     

Ph I-induced transfer of leaf and stripe rust-resistance genes from Aegilops triuncialis and Ae. geniculata to bread wheat

Leaf and stripe rusts are severe foliar diseases of bread wheat. Recently, chromosomes 5M^g from the related species Aegilops geniculata that confers resistance to both leaf and stripe rust and 5U^t from Ae. triuncialis conferring resistance to leaf rust have been transferred to bread wheat in the form of disomic DS5M^g(5D) and DS5U^t(5A) chromosome substitution lines. The objective of this study was to shorten the alien segments in these lines using Ph ^I-mediated, induced homoeologous recombination. Putativerecombinants were evaluated for their rust resistance, and by genomic in situ hybridization and microsatellite analyses. One agronomically useful wheat-Ae. geniculata recombinant resistant to leaf and stripe rust was identified that had only a small terminal segment of the 5M^gL arm transferred to the long arm of an unidentified wheat chromosome. This germplasm can be used directly in breeding programs. Only one leaf rust-resistant wheat-Ae. triuncialis recombinant, which consists of most of the complete 5U^t chromosome with a small terminal segment derived from 5AS, was identified. This germplasm will need further chromosome engineering before it can be used in wheat improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Wheat pre-breeding using wild progenitors

To facilitate the use of wheat wild relatives in conventional breedingprograms, a wheat pre-breeding activity started at ICARDA in 1994/1995season. Preliminary results of gene introgression from wild diploidprogenitors, Triticum urartu, T. baeoticum, Aegilops speltoides andAe. tauschii and tetraploid T. dicoccoides are described. Crosseswith wild diploid Triticum spp. yielded high variation in plant andspike morphology. Synthetic hexaploids were produced from crosses of alocal durum wheat landrace `Haurani' with two Ae. tauschiiaccessions. Both Ae. tauschii accessions carry hybrid necrosis allelesthat gave necrotic plant phenotypes in crosses with some bread wheats.Backcross progenies with agronomical desirable traits, i.e. high spikeproductivity, short plant stature, earliness, drought tolerance and highproductive tillering, were identified in crosses of durum wheat with wild Triticum spp. and in a cross of one of the hexaploid synthetics with alocally adapted bread wheat cv. `Cham 6'. Resistance to yellow rust wasfound in durum wheat crosses with the three wild Triticum spp. andAe. speltoides and leaf rust resistance was identified in crosses withT. baeoticum and Ae. speltoides. The results show that wheatimmediate progenitors may be a valuable and readily accessible source ofnew genetic diversity for wheat improvement.     

Genome-specific markers of tetraploid wheats and their putative diploid progenitor species

The objective of this study was to isolate genome‐specific markers from the genomes of tetraploid wheats and the putative donor diploid species on the basis of random amplified polymorphic DNA analysis followed by cross‐hybridization. Twenty different Triticum and Aegilops species and accessions were analysed by polymerase chain reaction (PCR) using 30 random primers. The polymorphic PCR fragments were then isolated, labelled and used in cross‐hybridization screenings. The hybridization results established that one marker was specific to the Ae. speltoides S genome, two to the A genome, one to the B genome and five to the G genomes of polyploid species (and to the genomes of the corresponding progenitor species). Four markers were identified that were specific to both the B and G genomes. Analysis of the Triticum and Aegilops species and accessions supported the notion that Ae. speltoides is more closely related to the B and G genomes of polyploid wheat species than were other members of the Sitopsis section. The data also indicated that the B and G genomes had originated from different accessions of Ae. speltoides.     

Origin of Sv Genome of Aegilops variabilis and Utilization of the Sv as Analyser of the S Genome of the Aegilops Species in the Sitopsis Section

Hybrids of Aegilops variabilis (2n = 28, UUS^vS^v) with the low-pairing type lines of Ae. longissima, Ae. bicornis, Ae. searsii, Ae. speltoides and Ae. umbellulata were obtained. This is the first report of successful hybridization between Ae. variabilis and Ae. searsii. Meiotic analysis of these hybrids showed that the U genome of Ae. variabilis donated by Ae. umbelltilata remained nearly unchanged and Ae. longissima is the possible donor species of the S genome to Ae. variabilis. But the S^v genome and the S¹ genome of Ae. longissima are not completely homologous and are structurally differentiated by at least one interchange. According to their levels of homology with S^v through the means of chromosome associations, the four genomes of Sitopsis section species can be classified as follow: S^v? S¹ > S^b > S^s > S. The meiotic behaviour of the hybrids between the wheat ‘Chinese Spring’ and the four Sitopsis section species studied has not supported the view that one of the four species is the donor of the B genome of wheat.     

Screening relatives of wheat for snow mold resistance and freeing tolerance

Four hundred and eighty one accessions of species and subspecies of genera Triticum and Aegilops were evaluated for resistance to speckled snow mold caused by Typhula ishikariensis and for freezing tolerance. All Triticum and seven of Aegilops species were severely affected by snow mold. Among surviving Aegilops spp., only the Ae. cylindrica accessions exhibited resistance similar to that of the most resistant winter wheat cultivar, PI 173438. After repeated screening of accessions of Ae. cylindrica, 12 accessions were identified as having resistance similar to that of PI 173438; eleven of those were considered more freezing tolerant than PI 173438, but less than wheat cv. Valjevskaya, the freezing tolerant check. Accessions of Ae. cylindrica with snow mold resistance exceeding that of PI 173438, and with freezing tolerance, are currently being screened. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic analysis of the Aegilops longissima 3S chromosome carrying the Pm13 resistance gene

The distal region of the short arm of chromosome 3S from Aegilopslongissima, which carries the powdery mildew resistance gene Pm13, was introgressed into common wheat. Due to suppression of recombination between this region and corresponding wheat homoeologous segments, a possible strategy to construct a genetic map around the Pm13 gene was based on crosses between a wheat addition line carrying the Ae.longissima 3S chromosome and the corresponding 3S addition lines of Ae.searsii and Ae. variabilis. The efficiency of this strategy was evaluated by scoring recombination frequencies inprogenies derived from these crosses. Recombination between 3S chromosomes fromAe. searsii and Ae. longissimawas very low, whereas 26.5% recombinant alien chromosomes were obtained from the cross involving the Ae. variabilisand Ae. longissima 3S addition lines. These data were used to construct a3S chromosome map that resulted largely colinear to the consensus map of the homoeologous group 3 of wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Origin and variation of gliadin proteins associated with pasta quality in durum wheat

Summary Studies were made of the presence and frequency of occurrence of gliadin bands 42 and 45 in three samples of Aegilops sharonensis Eig and 59 samples of wild tetraploid wheat (Triticum dicoccoides Korn.) from natural distributions of these species in Israel.Two samples of Ae. sharonensis possessed a band in position 45 and one possessed no bands corresponding to either band 45 or band 42. In T. dicoccoides, band 45 was either present or not and 42 was always absent. In its grassy and intermediate growth habit forms, (believed to be more primitieve than the cercal forms) band 45 appeared to be more frequent than in the cereal form.The presence of band 45 in the Ae. sharonensis, and its relatively high frequency in T. dicoccoides, populations from Mt. Hermon (likely to be relatively free from introgression from cultivated tetraploid wheat) indicate the likelihood of a primary origin of the allele coding for band 45. The absence of band 42 from all Ae. sharonensis and T. dicoccoides populations in this study, indicates a more recent evolutionary origin of the allele coding for this band, possibly arising as a mutation during the domestication of tetraploid wheat.The results have implications for breeding programmes in tetraploid wheat.     

Attempts to remove gametocidal genes co-transferred to common wheat with rust resistance from <Emphasis Type="Italic">Aegilops speltoides</Emphasis>

Rust resistance genes (introgressions S24 and S13) transferred to hexaploid wheat from two Aegilops speltoides accessions could not be used commercially due to associated gametocidal (Gc) genes. Crosses to wheat followed by rigorous selection for increased fertility were employed in an attempt to separate the unmapped S24 stem rust resistance from the Gc gene(s). However, improved fertility of the better selections could not be maintained in subsequent generations. Since the S13 introgression (leaf, stripe and stem rust resistances) mapped to chromosome 3A, allosyndetic pairing induction was used in an attempt to remove the Gc gene(s). This produced putative primary recombinants with improved fertility and plant type, the best of which had exchanged a small region of Ae. speltoides chromatin, yet was still associated with (reduced) Gc effects. This selection (04M127-3, which appears to have the Su1-Ph1 suppressor) was then crossed with wheat. Surprisingly, the 04M127-3 gametocidal effect differed drastically from that of the original introgression allowing the recovery of 35 recombinant, leaf rust resistant progeny. Microsatellite and DArT markers showed that each secondary recombinant had exchanged most of the Ae. speltoides chromatin. Although the data suggested that a complex multigenic interaction may govern the gametocidal response, preliminary indications are that the Gc effect had largely been removed and it now seems possible to completely separate the gametocidal genes from the S13 leaf rust resistance gene (here designated Lr66). The associated (S13) stripe rust and stem rust resistance genes were lost during recombination.     

The study of introgressive lines of Triticum aestivum × Aegilops speltoides by in situ and SSR analyses

Fluorescence in situ hybridization (FISH) with a genome‐specific repeat, Spelt1, and wheat simple sequence repeat (SSR) markers were used to analyse the chromosome constitution of two Triticum aestivum×Aegilops speltoides introgressive lines. The lines 170/98i and 178/98i carried one and two subtelomeric regions of Ae. speltoides (per haploid genome), respectively, marked by Spelt1 repeats according to FISH data. SSR analysis detected homoeologous substitution of wheat chromosome 7D with Ae. speltoides chromosome 7S in the lines 178/98i and 170/98i as well as the assumed terminal translocation in the short arm of chromosome 3A in the line 178/98i. Anthocyanin pigmentation of the coleoptiles was found in the lines 170/98i and 178/98i and resulted from the 7S (7D) substitution. It was demonstrated that Spelt1 could be effectively used for the rapid identification (without DNA isolation) of terminal translocations of T. aestivum×Ae. speltoides introgressive lines as well as for further analysis of the stability of the hybrid plants.     

Tan-Spot Resistance Screening of Aegilops Species

Two hundred and twelve accessions of 8 diploid and 10 polyploid species of Aegilops were evaluated for resistance to tan-spot disease of wheat, caused by Pyrenophora tritici-repentis (Died.) Drechs., under greenhouse conditions. One or more accessions of Ae. bicornis, Ae. biuncialis, Ae. Crassa, Ae. columnaris, Ae. cylindrica, Ae. speltoides, Ae. squarrosa. Ae. triaristata. Ae. triuncialis, and Ae. Ovata showed resistance following a 24-hour post-inoculation wet period. With a 36-hour wet period, diploids became only slightly or moderately susceptible and resistant polyploids became susceptible. A 48-hour wet period resulted in still greater susceptibility of both diploid and polyploid species.     

Microsatellite Marker Identification of a Triticum Aestivum -Aegilops Umbellulata Substitution Line with Powdery Mildew Resistance

Summary Aegilops umbellulata acc. Y39 and Triticum carthlicum acc. PS5, immune to many powdery mildew isolates, were crossed to make an amphidiploid line Am9. The powdery mildew resistance of Am9 was transferred to common wheat cultivar Laizhou953 by crossing and backcrossing. In this study, the origin of powdery mildew resistance in a BC₃F_4:5 population derived from a cross of Am9 and Laizhou953 was identified. Microsatellite markers analysis showed that markers Xgwm257, Xgwm296, and Xgwm319, co-segregated with the powdery mildew resistance, whereas markers Xgwm210, Xgwm388/140, Xgwm388/170 and Xgwm526 were related to susceptibility and linked to resistance in repulsion. Of three markers related to resistance, Xgwm257 and Xgwm319 were codominant, whereas Xgwm296 was dominant. All three markers were Ae. umbellulata-specific indicating that resistance in the test population originated from Ae. umbellulata acc. Y39. The chromosome location and mapping of these linked microsatellite markers, the chromosome numbers of derived BC₃F_4:6 families, and chromosome pairing in F₁ plants from a cross of a homozygous resistant BC₃F_4:5 plant and Laizhou953, showed that wheat chromosome 2B was substituted by Ae. umbellulata chromosome 2U. This is the first gene conferring powdery mildew resistance transferred to wheat from Ae. umbellulata, and it should be a novel resistance gene to powdery mildew. It was temporarily designated PmY39.The first two authors made equal contributions     

The Inheritance of Resistance to Septoria Glume Blotch III. The Wild Wheat Species Aegilops Jongissima

The genetics of resistance to Septoria glume blotch (caused by the pathogen Septoria nodorum Berk.) in the wild wheat species Ae. longissima was investigated. The resistance was characterized by two parameters measured on detached leaves — lesion size (LS) and length of latent period (LP), and by disease severity (DS) under field conditions. Generations F₁, F₂ and F₃, derived from a cross between two Ae. longissima accessions, were analyzed. The two parameters measured on detached leaves (LS and LP) were highly correlated, while DS was moderately correlated to both LS and LP. The mean LS and the mean LP of F₁ generation indicated considerable dominance for resistance in both parameters. The estimates of broad-sense and narrow-sense heritability were moderate for LS and LP (0.21—0.55). Narrow-sense heritability for DS was high (0.77). Estimates of the number of genes controlling each of the parameters (LS, LP, DS) were between 2.5—3.2. It is suggested that the resistance is controlled by three to four quantitative genes with a partial dominance of the alleles for resistance. Indications for genie interaction were found in LS and in LP. A model of inheritance containing complementation between dominant resistance-alleles is suggested. Highly resistant Ae. longissima accessions are recommended as sources of germplasm for improving the resistance of cultivated wheats to Septoria glume blotch. The possibility of using dominant alleles for resistance in hybrid cultivars is discussed.     

Characterisation and origin of rust and powdery mildew resistance genes in VPM1 wheat

Summary The expression of rust resistances conferred by closely linked genes derived from VPM1 varied with environmental conditions and with genetic backgrounds. Under low light and low temperature conditions seedlings carrying Yr17 showed susceptible responses. Stem rust and leaf rust resistance genes Sr38 and Lr37 tended to confer more resistance at 17±2° C than at normal temperatures above > 20° C. These studies supported the hypothesis that Yr17, Lr37 and Sr38 were derived from Aegilops ventricosa, whereas Pm4b was probably derived from T. persicum. Studies on certain addition lines and parental stocks indicated that wheat cytoplasm may enhance the expression of Sr38.     

Use of a synthetic hexaploid Triticum miguschovae for transfer of leaf rust resistance to common wheat

Summary Triticum miguschovae, a genome addition synthetic, was used as a source for transfer of leaf rust (Puccinia recondita tritici) resistance to common wheat. This synthetic, developed from two wild species Triticum militinae and Aegilops squarrosa, proves a valuable donor of the genes for leaf rust resistance. Leaf rust resistance was transferred from T. miguschovae by both dominant and recessive genes. Stable lines phenotypically similar to their recurrent parents Kavkaz and Bezostaya 1 but differing from them in a high level of leaf rust resistance were obtained. The genes for resistance in 3 selected lines differed from each other and from the known effective genes Lr9, Lr19, and Lr24. The resistance of one of them (line 1229) is controlled by two complementary interacting genes located on chromosome 7B and 1D was revealed by monosomic analysis.     

The Inheritance of Resistance to Septoria Glume Blotch II. The Wild Wheat Species Aegilops speltoides

The inheritance of resistance in the wild wheat species Ae. speltoides L. to Septoria glume blotch (Septoria nodorum Berk.) was investigated. It was characterized by two parameters measured on detached leaves, namely lesion size (LS) and length of latent period (LP), and by the severity of the disease under field conditions (DS). The genetic analysis was based on the F₁ and F₂ generations of a 4 × 4 diallel cross between four Ae. speltoides accessions showing differential resistance to the pathogen. The three parameters of resistance were highly correlated. Considerable heterosis for resistance was found for each of the parameters in most of the diallel combinations. The estimates of broad-sense heritability were moderately high (0.50 — LS, 0.60 — LP, 0.52 — DS), while the estimates of narrow-sense heritability were low (0.16 for LS, 0.20 for LP, 0.25 for DS). There were no indications for genie interaction. It is suggested that the resistance is controlled by at least two genes with over-dominance of the alleles for higher resistance. Highly resistant Ae. speltoides accessions are a potential source of germplasm for improving the resistance of cultivated wheats to Septoria glume blotch. The possibility of using dominant alleles for resistance in hybrid cultivars is discussed.     

Molecular mapping of genes determining hairy leaf character in common wheat with respect to other species of the Triticeae

Two major genes controlling leaf pubescence were mapped on chromosomes 4BL (Hl1) and 7BS (Hl2 ^Aesp ) in wheat (Saratovskaya 29) and a wheat/Aegilops introgression line (102/00^I), respectively, together with quantitative trait loci (QTLs) determining hairiness of the leaf margin (QHl.ipk-4B, QHl.ipk-4D) and auricle (QPa.ipk-4B, QPa.ipk-4D) on the long arms of chromosomes 4B and 4D, respectively. The QTLs on chromosome 4D were contributed by a synthetic wheat and, therefore, originated from Aegilops tauschii. The homoeologous group 4 wheat/A. tauschii genes/QTLs detected in the present study were aligned with the barley pubescence genes Hln/Hsh and Hs _b and the hairy peduncle rye gene Hp1. The locus seems to be pleiotropically responsible for the pubescence of different plant organs in different species of the Triticeae. Another homoeologous series may be present on the short arms of the homoeologous group 7 chromosomes, based on the results of an allelic test cross between the Chinese local cultivar Hong-mang-mai carrying Hl2 and the wheat/Aegilops speltoides introgression line 102/00^I.