Resistance to wheat leaf rust in Aegilops tauschii Coss. and inheritance of resistance in hexaploid wheat

A collection of 164 Aegilops tauschii accessions, obtained from Gatersleben, Germany, was screened for reaction to leaf rust under controlled greenhouse conditions. We have also evaluated a selection of synthetic hexaploid wheats, produced by hybridizing Ae. tauschii with tetraploid durum wheats, as well as the first and second generation of hybrids between some of these resistant synthetic hexaploid wheats and susceptible Triticum aestivum cultivars. Eighteen (11%) accessions of Ae. tauschii were resistant to leaf rust among which 1 was immune, 13 were highly resistant and 4 were moderately resistant. Six of the synthetic hexaploid wheats expressed a high level of leaf rust resistance while four exhibited either a reduced or complete susceptibility compared to their corresponding diploid parent. This suppression of resistance at the hexaploid level suggests the presence of suppressor genes in the A and/or B genomes of the T. turgidum parent. Inheritance of leaf rust resistance from the intercrosses with susceptible bread wheats revealed that resistance was dominant over susceptibility. Leaf rust resistance from the three synthetics (syn 101, syn 701 and syn 901) was effectively transmitted as a single dominant gene and one synthetic (syn 301) possessed two different dominant genes for resistance.     

Distribution and diversity of Aegilops tauschii in Iran

The wide morphological variation of Aegilops tauschii has led to the distinction of different subspecies; a typical ssp. tauschii and a second ssp. strangulata. However some researchers pointed out the existance of the intermediate form among morphologically distinguished subspecies. Distribution, diversity and the relationship between different subspecies and the intermediate form were evaluated in the Iranian Ae. tauschii collection. This collection was classified to 15 different populations according to morphological similarities and the collecting origin of accessions. The highest variation was found in tauschii population of Golestan followed by tauschii populations of Gilan and Ardebill, whereas the lowest variation was observed in tauschii populations of central Iran. Two discriminant functions suggested that the length of rachis node and spikelet glume, particularly, the length/width ratios of these traits had the highest impact on identification of different forms. Mahalanobis distances (D ² ) between the two subspecies along with intermediate form on the multidimensional scaling plot showed that the intermediate form is more similar to ssp. tauschii than ssp. strangulata. Although, the diversity within the ssp. strangulata was not very high, it widely affected the diversity of Iranian accessions of Ae. tauschii through continues crossing with the more diversed subspecies, tauschii, during thousands of years. This fact had lead to expansion of its distribution from its origin to Northern Khorasan, Northern Semnan and Eastern Ardebill by producing the intermediate form.     

Novel and ancient HMW glutenin genes from Aegilops tauschii and their phylogenetic positions

A pair of novel high-molecular-weight glutenin subunits (HMW-GS) 1Dx3.1^t and 1Dy11*^t were revealed and characterized from Aegilops tauschii Coss. subspecies tauschii accession AS60. SDS-PAGE band of 1Dx3.1^t was between those of 1Dx2 and 1Dx3, while 1Dy11*^t was between 1Dy11 and 1Dy12. The lengths of 1Dx3.1 ^t and 1Dy11* ^t were 2,514?bp and 1,968?bp, encoding 836 and 654 amino acid residues, respectively. Their authenticity was confirmed by successful expression of the coding regions in Escherichia coli. Network analysis indicated that 1Dx3.1 ^t together with other five rare alleles only detected in Asia common wheat populations represented the ancestral sequences in Glu-D1 locus. Neighbor-joining tree analysis of previously cloned x-type and y-type alleles in the Glu-D1 locus supported the hypothesis that more than one Ae. tauschii genotypes were involved in the origin of hexaploid wheat and that different Ae. tauschii accessions contributed the D genome to common wheat and Ae. cylindrical Host, respectively. An Ae. tauschii accession with 1Dx3.1 ^t or a closely related allele probably have involved in the origin of common wheat. Since accession AS60 used in this study belonged to typical ssp. tauschii, present results suggested the possibility that ssp. tauschii was involved in the evolution of common wheat.     

Evaluation for resistance to Pyrenophora tritici-repentis in Aegilops tauschii Coss. and synthetic hexaploid wheat amphiploids

Summary A total of 59 diploid Aegilops tauschii Coss. (syn. Aegilops sguarrosa auct. non L.) and 39 synthetic hexaploid wheat accessions were evaluated for reaction to Pyrenophora tritici-repentis (Died.) Drechs. in a controlled environment, and classified using a disease rating system based on lesion type. 27 Ae. tauschii and 20 synthetic wheats were found to be resistant to tan spot disease. The overall mean disease ratings of Ae. tauschii and the synthetic wheat lines scored on a scale of 1 (resistant) to 5 (susceptible) were 1.80 and 2.38, respectively. Synthetic wheats generally showed a decrease in resistance, although several lines of synthetic wheat expressed a higher resistance than the diploid parents. Five synthetic wheat lines exhibited higher resistance than the standard resistant common wheat cultivar Red Chief.     

Amphidiploids between tetraploid wheat and Aegilops sharonensis Eig exhibit variations in high-molecular-weight glutenin subunits

Wide crossing is a useful method for transferring traits of interest from wild Triticeae species to wheat. In this study, Aegilops sharonensis was successfully hybridized to three tetraploid wheat species with crossability varying from 0.86 to 1.76 %. F1 hybrids were verified by their display of intermediate characteristics in morphology, cytology, and sodium dodecyl sulfate–polyacrylamide gel electrophoresis profiles. The exogenous x- and y-type subunits of high-molecular-weight glutenin subunits of Ae. sharonensis were expressed in F1 hybrid seeds and showed identical electrophoretic mobilities to those originally present in Ae. sharonensis. In F2 hybrids, the x-type subunit migrated faster, indicating that a decrease in molecular mass had occurred. Cloning of the open reading frames of subunit variants revealed that the x-type subunit had protein sequence variations, including single residue mutations, and insertions and deletions involving one or more repeat motifs. The two deletions causing the size reduction of the exogenous x-subunits of Ae. sharonensis in the F2 seeds were most likely the result of slip-mismatching.     

Powdery mildew resistance in Aegilops tauschii coss. and synthetic hexaploid wheats

Summary A collection of 400 Ae. tauschii (syn. Ae. squarrosa) Coss. accessions were screened for powdery mildew resistance based on the response patterns of 13 wheat cultivars/lines possessing major resistance genes to nine differential mildew isolates. 106 accessions showed complete resistance to all isolates, and 174 accessions revealed isolate-specific resistance, among which were 40 accessions exhibiting an identical response pattern as wheat cultivar Ulka/^*8Cc which is known to possess resistance gene Pm2. Expression of both complete and isolate-specific resistance from Ae. tauschii was observed in some synthetic hexaploid wheats derived from four mildew susceptible T. durum Desf. parents, each crossed with five to 38 resistant diploid Ae. tauschii accessions. Synthetic amphiploids involving different combinations of T. durum and Ae. tauschii generally showed a decrease in resistance compared with that expressed by the Ae. tauschii parental lines.     

Evaluation of shape Aegilops tauschii Coss for resistance to physiological strains CYR30 and CYR31 of wheat stripe rust in China

Stripe rust is one of the most serious diseases of wheat in China. Two new stripe rust physiological strains CYR₃₀ (Inter. name: 175 E 191) and CYR₃₁ (Inter. name: 239 E 175) have become the dominant and epidemic physiological strains since 1994 in China. Resistance to these strains of 48 Ae. tauschii accessions was evaluated at the seedling and adult stages. Out of 48, 28 accessions displayed seedling and adult resistance. All of the Chinese Ae. tauschii accessions were susceptible at seedling and adult stages. Seedling resistance was highly related to adult resistance in Ae. tauschii. The new genetic resources of resistance to CYR₃₀ and CYR₃₁ could be incorporated into commercial wheat varieties for wheat resistance breeding by direct hybridization.     

Multivariate analysis of genetic variation in Aegilops tauschii from the world germplasm collection

A study of Aegilops tauschii subspecies constitution was undertaken. The data on allozyme and morphologic variation among 308 plants from 154 accessions were used for multivariate analysis. ACPH1 and (glume width)/(rachis segment width) ratio were found to be reliable criteria to distinguish between sspp. tauschii and strangulata.     

Aegilops tauschii Coss.: allelic variation of enzyme-encoding genes and ecological differentiation of the species

Three hundred and seven accessions of Aegilops tauschii Coss., including 160 of subsp. tauschii and 147 of subsp. strangulata, representing all the species range—from Turkey to Kirgizstan, were analyzed electrophoretically. Twenty polymorphic enzyme-encoding loci were studied, 10 of which were essentially polymorphic in Ae. tauschii. Climatic data for each of the 307 Ae. tauschii habitats were taken from WORLDCLIM database of computer system ArcGIS. Forty-nine climatic parameters were considered: precipitation, minimum, mean and maximum temperatures for each month, and also the total annual level of precipitation. The data were analyzed with multivariate statistical methods, such as Principal Components Analysis (PCA), Multiple Correspondence Analysis (MCA) and Two-Block Partial Least Squares. Variability of climatic conditions among Ae. tauschii habitats is reflected by the two approximately orthogonal “vectors”. The “first vector” is mostly determined by negative impact of precipitation and minimum temperatures during winter. The “second vector” is mostly determined by negative impact of maximum temperatures during summer, and positive impact of precipitation during late spring and summer. Aegilops tauschii is essentially variable along the “second vector”, and especially high level of variation is characteristic for subsp. tauschii. This variation reflects that Ae. tauschii is very tolerable to the climatic variation during summer season. Aegilops tauschii subsp. strangulata is also characterized by the high level of variation along the “first vector”. Moreover, all the habitats of subsp. strangulata fall into the two distinct separate clusters: the habitats in Precaspian Iran, which have the highest minimum temperatures in winter,—and all the other habitats. In the plot of the first two factors of PCA, the “cluster of Precaspian Iran” can be further divided into “Western Precaspian Iran (WPI)”, having relatively higher level of annual rainfall, and relatively dryer “Eastern Precaspian Iran (EPI)”. This three groups of subsp. strangulata accessions, from WPI, EPI and other areas, are also distinctly differed in enzyme-encoding genes allelic variation, as revealed on the plot of the first two axes of MCA. In contrast to subsp. strangulata, the level of variation of subsp. tauschii along the “first vector” is rather low. It was pointed out that variation along “the first vector” reflects adaptive intraspecies divergence of Ae. tauschii: its subspecies strangulata “prefers” the habitats of seaside climate, with warm and moist winter; while subsp. tauschii mostly occupies the habitats with rather continental climate, with relatively cold and dry winter. Allelic variation of enzyme-encoding genes Acph1, Ak, Est2, Est5, Got1, Got2, and Got3 correlate with climate along “the first vector”. Apparently, polymorphism of these loci were involved into the process of Ae. tauschii intraspecies adaptive divergence. Allelic variation of Cat2 and Fdp loci correspond to climatic variation along “the second vector” in subsp. tauschii. Therefore Cat2 and Fdp are likely to be among the genes which polymorphism “helped” subsp. tauschii to succeed in vast geographical expansion far to the east from Caspian Sea.     

Chloroplast DNA non-coding sequences variation in Aegilops tauschii Coss.: evolutionary history of the species

Sequences of four chloroplast DNA non-coding regions, about 3,000?bp in total, were analysed in 112 Aegilops tauschii accessions, 56 of ssp. tauschii and 56 of ssp. strangulata, representing all of the species range. One inversion, 8 insertions/deletions, 18 base pair substitutions and 5 microsatellite loci were found. The data revealed that Ae. tauschii originated in Caucasia. Neither of the two Ae. tauschii subspecies was an ancestor to one another. Aegilops tauschii divided into ssp. tauschii and ssp. strangulata at the very beginning of its existence as a species. Subspecies tauschii was the first to start geographic expansion and relatively rapidly occupied a vast area from Caucasia—eastward up to central Tien Shan and western Himalayas. In contrast to ssp. tauschii, geographic spread of ssp. strangulata was a complicated, multi-stage and slow process. At the beginning of ssp. strangulata evolutionary history its major phylogenetic lineage for a lengthy time span had existed as a small isolated population. Several forms of ssp. strangulata, better adapted to relatively moister and cooler habitats, had originated. Each of these forms has gradually forced out ssp. tauschii from some part of its area in the west, up to central Kopet-Dag.     

Geographic patterns of Got1, Got2, Got3 and Est2 enzyme-encoding genes allelic variation in Aegilops tauschii Coss.

Geographic patterns of Got1, Got2, Got3 and Est2 enzyme-encoding genes allelic variation were investigated among 322 accessions of Aegilops tauschii Coss., 161 accessions of ssp. tauschii and 161 accessions of ssp. strangulata, representing all the species area. It was found that: (1) in the two ecologically different subspecies, ssp. tauschii and ssp. strangulata, the patterns of allelic variation of the four genes differ greatly; (2) the same allozymes have originated several times independently in different Ae. tauschii local populations; (3) allelic variation of Got1, Got3 and Est2 in ssp. strangulata corresponds to climatic conditions. The data obtained reflected that Ae. tauschii has been inhabiting its area from ancient times; and allelic variation patterns of Got1, Got2, Got3, Est2 loci were mostly formed by natural selection. Further investigations of these loci with molecular genetic methods are prospective for understanding the peculiarities of Ae. tauschii evolution.     

Single-copy nuclear PolA1 gene sheds light on the origin of S genome with relationships to B and G genomes of polyploid wheat species

PolA1, a single-copy nuclear gene encoding the largest subunit of RNA polymerase I, comprises highly polymorphic intron 19 and nucleotide tag (Ntag) sequences. We analyzed these sequences in 42 accessions, which differed in ploidy, of Triticum–Aegilops and Hordeum species. The lengths of the intron 19 sequences were ca. 110?bp long in Triticum–Aegilops species, except in four Sitopsis species, Ae. longissima, Ae. searsii, Ae. sharonensis, Ae. speltoides, which had introns similar in length to those of Hordeum species, i.e., ca. 240?bp long. Phylogenetic analyses of the Ntag sequences showed that the four Sitopsis and remaining Triticum–Aegilops species were classified into two discrete Hordeum and Triticum clades, respectively. The A and D genome-specific Ntag sequences of polyploid wheats were highly homologous with those of T. urartu and Ae. tauschii, respectively. In Ae. bicornis, another Sitopsis species, two accessions had the short intron 19 and Triticum–type Ntag sequence, which were highly homologous with those of the B genome in polyploid wheats, whereas one accession contained the long intron 19 and Hordeum–type Ntag sequences. In contrast, partial sequence analyses revealed that the three accessions of Ae. bicornis shared highly homology to single-copy DMC1 and EF-G genes. The discrepancy between these results indicates that the Sitopsis species were probably established by hybrid speciation including ancient introgressive hybridization between progenitors of Triticum–Aegilops and Hordeum. Although many researchers have proposed Ae. speltoides as a candidate for the B genome donor, our data suggest the existence of diploid B genome species in the past that were responsible for the origin of both polyploid wheats and Sitopsis species, including Ae. speltoides.     

Morphological and isozyme variation in genebank accessionsof Aegilops umbellulata Zhuk., a wild relative of wheat

Morphological variation and genetic variation at 15 enzyme lociwere studied in genebank accessions of Aegilopsumbellulata Zhuk., the diploid genome donor to all thepolyploid species of the section Aegilops ofAegilops. Accessions from the Greek Islands hadshorter spikes with smaller number of spikelets and smaller number ofawns on the empty glume. The number of alleles per locus (A= 2.01) and the proportion of polymorphic loci (P= 0.627) were similar to those of the other species ofAegilops so far reported. Genetic distancescalculated from isozyme variations among five regions revealed thataccessions from the Greek Islands are more distantly related to theother four continental regions, Iran and Iraq, Southeast Turkey,Central Turkey and South and West Turkey. The present resultsindicated the importance of collection covering the whole range ofgeographical distribution to capture the genetic variation present inAe. umbellulata.     

Phenotypic responses of the wild wheat relative Aegilops geniculata Roth and wheat (Triticum durum Desf.) to experimentally imposed salt stress

In order to study the salt effect on the wild wheat and durum wheat, three accessions of Aegilops geniculata L. from Ain zana, Zaghouan, and Sbitla and one variety of wheat (Triticum durum L.) have been grown in the INRAT green house and treated with different salt concentrations. The morphological, phenological and yield characters have been measured for each plant and analyzed using SAS software. This study has shown a high degree of variation of these characters mainly related to geographical origin. It was observed also that Sbitla accession was less affected by the imposed salt stress than all the others while Ain zana was the most affected one.     

Conservation of the genetic variation of Triticum tauschii (Coss.) Schmalh. (Aegilops squarrosa auct. non L.) in synthetic hexaploid wheats (T. turgidum L. s.lat. x T. tauschii; 2n=6x=42, AABBDD) and its potential utilization for wheat improvement

Triticum tauschii (Coss.) Schmalh. (Aegilops squarrosa auct. non L., 2n=2x=14, DD genome) with its diverse range of accessions and distribution provides a unique opportunity for exploiting novel genetic variability for wheat (T. aestivum L.) improvement associated with biotic/abiotic stress factors. From our working collection of 490 T. tauschii accessions we have so far produced 430 different synthetic hexaploids (2n=6x=42, AABBDD) resulting from the chromosome doubling of Triticum turgidum L. s. lat. x T. tauschii F₁ hybrids (each synthetic involving a different T. tauschii accession). We present here our results on hybrid production, plantlet regeneration, cytology, colchicine induced doubling of the 2n=3x=21 chromosome F1 hybrids, seed increase of the doubled progeny and screening for a biotic stress; Cochliobolus sativus Ito and Kuribay (syn. Helminthosporium sativum Pamm. King and Bakke); of 250 of these synthetic hexaploid (2n=6x=42) amphiploids. Application of the direct crossing methodology involving susceptible T. aestivum cultivars with resistant T. tauschii accessions is also alluded to.     

The climate of the zone of origin of Mediterranean durum wheat (Triticum durum Desf.) landraces affects their agronomic performance

The genetic diversity of durum wheat (Triticum durum Desf.) is held by landraces, which are generally considered to be endemic to a particular region to which they are well adapted. To evaluate the effect of climate in the countries of origin on their agronomic performance, 172 durum wheat landraces from 21 Mediterranean countries were grown in northeastern Spain. Average long-term climatic data of the main wheat-growing areas in each country of origin allowed us to identify four climatic zones in the Mediterranean Basin, steadily varying from warm and dry to cool and wet. The phenology, biomass, and yield of landraces were affected by the climatic zone of origin. The climatic zone accounted for 32.8, 28.3 and 14.5 % of variance for days to anthesis, plant height, and grain filling rate, respectively. The number of days to heading and anthesis steadily increased when moving from the warmest and driest zone of origin to the coldest and wettest one. Landraces collected in the warmest and driest zone had a smaller biomass, a lower chlorophyll content in the flag leaf, more fertile tillers, spikes and grains m^?2, a lower grain filling rate, lighter grains, and lower yields than those originated in colder and wetter zones. Landraces collected in countries with high solar radiation showed a shorter cycle until anthesis and smaller height and biomass accumulation, while higher temperatures after anthesis resulted in more tillers and spikes. Landraces from countries with high potential evapotranspiration during grain filling had significantly lower grain filling rates and grain weight.     

Role of chemical constituents of wheat straw and pig slurry on their decomposition in soil

Carbon mineralization and microbial biomass content of wheat straw (WS), pig slurry (PS) and their mixture (WSPS), either intact or with extraction of soluble substances (–SS) or soluble substances plus hemicellulose (–SSH), added to soil, were monitored over 230 days in a laboratory incubation experiment. The WSPS showed a CO₂ release of up to 23% above that predicted by summing the CO₂ evolved from WS and PS. Of the several kinetic models tested to describe the mineralization process, a double exponential model best described the C mineralization of all the materials, both intact and with extractions. The extraction of the labile substances from WS, PS and WSPS lowered the values of the rapidly mineralizable C and of the amount of microbial biomass. The organic fraction of WS was found to be almost completely represented by mineralizable carbon, while PS and WSPS showed only 62% of mineralizable carbon. In spite of this, after 8 months, about half of the initial amount of the organic C in the intact residues still remained unmineralized. Received: 29 October 1996     

Sources of resistance to Hessian fly (Diptera: Cecidomyiidae) in Syria identified among Aegilops species and synthetic derived bread wheat lines

The Hessian fly, Mayetiola destructor (Say), is a major pest of wheat in North Africa, southern Europe, North America, and northern Kazakhstan. It is believed this pest (like wheat) originated in West Asia. The Syrian Hessian fly biotype has been found to be the most virulent worldwide, and has been used at the International Center for Agricultural Research in the Dry Areas (ICARDA) for screening wheat and its wild relatives to identify new sources of resistance. The screening was conducted in an insect rearing room set at 20°C and 70% RH using a Hessian fly population collected from Lattakia region, Syria. The experimental design was a randomized complete block with four replications. ‘Nasma’ (bread wheat) and ‘Cando’ (durum wheat) were used as susceptible and resistant checks, respectively. A total of 623 lines/accessions of wheat and its wild relatives (Aegilops and Triticum) were evaluated. Twenty-nine Aegilops accessions and four synthetic derived bread wheat lines were found resistant. The presence of dead first instars confirmed the resistance reaction and also showed that antibiosis is the major mechanism of resistance in these materials. These sources of resistance are used in ICARDA’s wheat breeding programs for the development of Hessian fly-resistant germplasm/varieties.