Heterochromatin differentiation and phylogenetic relationship of the A genomes in diploid and polyploid wheats

Summary Heterochromatin differentiation, including band size, sites, and Giemsa staining intensity, was analyzed by the HKG (HCl-KOH-Giemsa) banding technique in the A genomes of 21 diploid (Triticum urartu, T. boeoticum and T. monococcum), 13 tetraploid (T. araraticum, T. timopheevi, T. dicoccoides and T. turgidum var. Dicoccon, Polonicum), and 7 cultivars of hexaploid (T. aestivum) wheats from different germplasm collections. Among wild and cultivated diploid taxa, heterochromatin was located mainly at centromeric regions, but the size and staining intensity were distinct and some accessions' genomes had interstitial and telomeric bands. Among wild and cultivated polyploid wheats, heterochromatin exhibited bifurcated differentiation. Heterochromatinization occurred in chromosomes 4A^t and 7A^t and in smaller amounts in 2A^t, 3A^t, 5A^t, and 6A^t within the genomes of the tetraploid Timopheevi group (T. araraticum, and T. timopheevi) and vice versa within those of the Emmer group (T. dicoccoides and T. turgidum). Similar divergence patterns occurred among chromosome 4A^a and 7A^a of cultivars of hexaploid wheat (T. aestivum). These dynamic processes could be related to geographic distribution and to natural and artifical selection. Comparison of the A genomes of diploid wheats with those of polyploid wheats shows that the A genomes in existing diploid wheats could not be the direct donors of those in polyploid wheats, but that the extant taxa of diploids and polyploids probably have a common origin and share a common A-genomelike ancestor.Contribution of the College of Agricultural Sciences, Texas Tech Univ. Journal No. T-4-233.     

NADP-dependent aromatic alcohol dehydrogenase in polyploid wheats and their diploid relatives. On the origin and phylogeny of polyploid wheats

Summary The three major isoenzymes of the NADP-dependent aromatic alcohol dehydrogenase (ADH-B), distinguished in polyploid wheats by means of polyacrylamide gel electrophoresis, are shown to be coded by homoeoalleles of the locus Adh-2 on short arms of chromosomes of the fifth homoeologous group. Essentially codominant expression of the Adh-2 homoeolleles of composite genomes was observed in young seedlings of hexaploid wheats (T. aestivum s.l.) and tetraploid wheats of the emmer group (T. turgidum s.l.), whereas only the isoenzyme characteristic of the A genome is present in the seedlings of the timopheevii-group tetraploids (T. timopheevii s.str. and T. araraticum).The slowest-moving B³ isoenzyme of polyploid wheats, coded by the homoeoallele of the B genome, is characteristic of the diploid species Aegilops speltoides S.l., including both its awned and awnless forms, but was not encountered in Ae. bicornis, Ae. sharonensis and Ae. longissima. The last two diploids, as well as Ae. tauschii, Ae. caudata, Triticum monococcum s.str., T. boeoticum s.l. (incl. T. thaoudar) and T. urartu all shared a common isoenzyme coinciding electrophoretically with the band B² controlled by the A and D genome homoeoalleles in polyploid wheats. Ae. bicomis is characterized by the slowest isoenzyme, B⁴, not found in wheats and in the other diploid Aegilops species studied.Two electrophoretic variants of ADH-B, B¹ and B², considered to be alloenzymes of the A genome homoeoallele, were observed in T. dicoccoides, T. dicoccon, T. turgidum. s.str. and T. spelta, whereas B² was characteristic of T. timopheevii s.l. and only B¹ was found in the remaining taxa of polyploid wheats. The isoenzyme B¹, not encountered among diploid species, is considered to be a mutational derivative which arose on the tetraploid level from its more ancestral form B² characteristic of diploid wheats.The implication of the ADH-B isoenzyme data to the problems of wheat phylogeny and gene evolution is discussed.     

Microsatellite mapping of <Emphasis Type="Italic">Ae. speltoides</Emphasis> and map-based comparative analysis of the S,G, and B genomes of Triticeae species

The first microsatellite linkage map of Ae. speltoides Tausch (2n = 2x = 14, SS), which is a wild species with a genome closely related to the B and G genomes of polyploid wheats, was developed based on two F₂ mapping populations using microsatellite (SSR) markers from Ae. speltoides, wheat genomic SSRs (g-SSRs) and EST-derived SSRs. A total of 144 different microsatellite loci were mapped in the Ae. speltoides genome. The transferability of the SSRs markers between the related S, B, and G genomes allowed possible integration of new markers into the T. timopheevii G genome chromosomal maps and map-based comparisons. Thirty-one new microsatellite loci assigned to the genetic framework of the T. timopheevii G genome maps were composed of wheat g-SSR (genomic SSR) markers. Most of the used Ae. speltoides SSRs were mapped onto chromosomes of the G genome supporting a close relationship between the G and S genomes. Comparative microsatellite mapping of the S, B, and G genomes demonstrated colinearity between the chromosomes within homoeologous groups, except for intergenomic T6A^tS.1G, T4AL.5AL.7BS translocations. A translocation between chromosomes 2 and 6 that is present in the T. aestivum B genome was found in neither Ae. speltoides nor in T. timopheevii. Although the marker order was generally conserved among the B, S, and G genomes, the total length of the Ae. speltoides chromosomal maps and the genetic distances between homoeologous loci located in the proximal regions of the S genome chromosomes were reduced compared with the B, and G genome chromosomes.     

Chromosome substitutions of Triticum timopheevii in common wheat and some observations on the evolution of polyploid wheat species

Whether the two tetraploid wheat species, the well known Triticum turgidum L. (macaroni wheat, AABB genomes) and the obscure T. timopheevii Zhuk. (A^tA^tGG), have monophyletic or diphyletic origin from the same or different diploid species presents an interesting evolutionary problem. Moreover, T. timopheevii and its wild form T. araraticum are an important genetic resource for macaroni and bread-wheat improvement. To study these objectives, the substitution and genetic compensation abilities of individual T. timopheevii chromosomes for missing chromosomes of T. aestivum Chinese Spring (AABBDD) were analyzed. Chinese Spring aneuploids (nullisomic-tetrasomics) were crossed with a T. timopheevii x Aegilops tauschii amphiploid to isolate T. timopheevii chromosomes in a monosomic condition. The F₁ hybrids were backcrossed one to four times to Chinese Spring aneuploids without selection for the T. timopheevii chromosome of interest. While spontaneous substitutions involving all A^t- and G-genome chromosomes were identified, the targeted T. timopheevii chromosome was not always recovered. Lines with spontaneous substitutions from T. timopheevii were chosen for further backcrossing. Six T. timopheevii chromosome substitutions were isolated: 6A^t (6A), 2G (2B), 3G (3B), 4G (4B), 5G (5B) and 6G (6B). The substitution lines had normal morphology and fertility. The 6A^t of T. timopheevii was involved in a translocation with chromosome 1G, resulting in the transfer of the group-1 gliadin locus to 6A^t. Chromosome 2G substituted for 2B at a frequency higher than expected and may carry putative homoeoalleles of gametocidal genes present on group-2 chromosomes of several alien species. Our data indicate a common origin for tetraploid wheat species, but from separate hybridization events because of the presence of a different spectrum of intergenomic translocations.     

New 18S·26S ribosomal RNA gene loci: chromosomal landmarks for the evolution of polyploid wheats

Three new 18S·26S rRNA gene loci were identified in common wheat by sequential N-banding and in situ hybridization (ISH) analysis. Locus Nor-A7 is located at the terminal area of the long arm of 5A in both diploid and polyploid wheats. Locus Nor-B6 is located in N-band 1BL2.5 of the long arm of chromosome 1B in Triticum turgidum and Triticum aestivum. ISH sites, similar to Nor-B6, were also detected on the long arms of chromosomes 1G in Triticum timopheevii and 1S in Aegilops speltoides, but their locations on the chromosomes were different from that of Nor-B6, indicating possible chromosome rearrangements in 1GL and 1BL during evolution. The third new locus, Nor-D8, was only found on the short arm of chromosome 3D in the common wheat Wichita. The loss of rRNA gene locus Nor-A3 and gain of repetitive DNA sequence pSc119 on the terminal part of 5AS suggest a structural modification of 5AS. Comparative studies of the location of the 18S·26S rRNA gene loci in polyploid wheats and putative A and B (G) genome progenitor species support the idea that: (1) Triticum monococcum subsp. urartu is the donor of both the A and A^t genome of polyploid wheats. (2) Ae. speltoides is closer to the B and G genome of polyploid wheats than Aegilops longissima and is the most probable progenitor of these two genomes.     

Wheat genome structure: translocations during the course of polyploidization

The genomic organization of Triticum timopheevii (2n=28, A^tA^tGG) was compared with hexaploid wheat T. aestivum (2n=42, AABBDD) by comparative mapping using microsatellites derived from bread wheat. Genetic maps for the two crosses T. timopheevii var. timopheevii × T. timopheevii var. typica and T. timopheevii K-38555×T. militinae were constructed. On the first population, 121 loci were mapped, and on the second population 103 loci. The transferability of the wheat markers to T. timopheevii was generally better for the A genome-specific markers (76–78% produced amplification products; 26 and 29% were polymorphic) than for B genome-specific markers (54% produced amplification products; 14 and 16% were polymorphic). Of the D genome-specific markers, one third produced amplification products in T. timopheevii, but only 5 and 2% were polymorphic in the corresponding mapping populations. The maps constructed confirmed the previously described translocation between chromosome arms 6A^tS and 1GS and revealed at least two yet unknown rearrangements on chromosomes 4A^t and 6A^t. The presence of other translocations and rearrangements between T. timopheevii and T. aestivum was demonstrated by a variety of markers mapping to nonhomoeologous positions.     

Single-copy gene fluorescence in situ hybridization and genome analysis: Acc-2 loci mark evolutionary chromosomal rearrangements in wheat

Fluorescence in situ hybridization (FISH) is a useful tool for physical mapping of chromosomes and studying evolutionary chromosome rearrangements. Here we report a robust method for single-copy gene FISH for wheat. FISH probes were developed from cDNA of cytosolic acetyl-CoA carboxylase (ACCase) gene (Acc-2) and mapped on chromosomes of bread wheat, Triticum aestivum L. (2n?=?6x?=?42, AABBDD), and related diploid and tetraploid species. Another nine full-length (FL) cDNA FISH probes were mapped and used to identify chromosomes of wheat species. The Acc-2 probe was detected on the long arms of each of the homoeologous group 3 chromosomes (3A, 3B, and 3D), on 5DL and 4AL of bread wheat, and on homoeologous and nonhomoeologous chromosomes of other species. In the species tested, FISH detected more Acc-2 gene or pseudogene sites than previously found by PCR and Southern hybridization analyses and showed presence/absence polymorphism of Acc-2 sequences. FISH with the Acc-2 probe revealed the 4A–5A translocation, shared by several related diploid and polyploid species and inherited from an ancestral A-genome species, and the T. timopheevii-specific 4A^t–3A^t translocation.     

Chromosome synteny of the a genome of two evolutionary wheat lines

In order to estimate synteny between A^t and A polyploid wheat genomes belonging to different evolutionary lines (Timopheevi and Emmer), saturation of chromosome maps of Triticum timopheevii A^t genome by molecular markers has been conducted. Totally, 179 EST-SSR and 48 genomic SSR-markers have been used with the following integration of 13 and 7 markers correspondingly into chromosome maps of A^t genome. ESTSSR showed higher transferability and lower polymorphism than genomic SSR markers. The chromosome maps designed were compared to maps of homoeologous chromosome group of the T. aestivum A genome. No disturbances of colinearity, i.e., of the order of markers within the chromosome segments on which they had been previously mapped, were observed. According to the quantity assessment of markers amplifying in homoeologous chromosomes, the maximum divergence was detected in two groups (4A^t/4A and 3A^t/3A) among the seven chromosomes examined in the A ^t and A genomes. Comparison of molecular genetic mapping results with the published results of studying meiosis of F₁ hybrids and the frequency of chromosomes substitution in introgressive T. aestivum × T. timopheevii lines suggest that individual chromosomes of the A^t and A genomes evolve differently. Translocations were shown to introduce the major impact on the divergence of 4A^t/4A and 6A^t/6A chromosomes, while mutations of the primary DNA structure, on the divergence of homoeologous group 3 chromosomes. The level of reorganization of other chromosomes during the evolution in the A^t and A genomes was significantly lower.     

Cytogenetic investigation of Triticum timopheevii (Zhuk.) Zhuk. and related species using the C-banding technique

Triticum timopheevii and related species T. militinae (2n=28, A^tG) and T. zhukovskyi (2n=42, A^mA^tG), hybrids T. kiharae, T. miguschovae, the amphidiploid T. timopheevii x T. tauschii (all 2n=42, A^tGD), T. fungicidum (ABA^tG) and T. timonovum (2n=56, A^tA^tGG) were analyzed using the C-banding technique. Chromosomes of the A^m and A^t genomes in the karyotype of T. zhukovskyi differed in their C-banding pattern. Partial substitutions of A^t-genome chromosomes and a complete substitution of the G-genome chromosomes by homoeologous chromosomes of an unidentified tetraploid wheat species with an AB genome composition were found in the T. timonovum karyotype. A^t- and G-genome chromosomes in the karyotypes of all studied species had similar C-banding patterns and were characterized by a low level of polymorphism. The comparative stability of the A^t and G genomes is determined by the origin and specifity of cultivation of studied species.     

A reassessment of the origin of the polyploid wheats

The diploid species that donated the A and D genomes to the polyploid wheats have been recognized for some time. New evidence indicates that Triticum speltoides cannot be the B genome donor to T. turgidum or T. aestivum. T. speltoides is probably homologous to the G genome of T. timopheevii. The donor of the B genome to T. turgidum and T. aestivum is currently unrecognized.     

NAD-dependent aromatic alcohol dehydrogenase in wheats (Triticum L.) and goatgrasses (Aegilops L.): evolutionary genetics

Summary Evolutionary electrophoretic variation of a NAD-specific aromatic alcohol dehydrogenase, AADH-E, in wheat and goatgrass species is described and discussed in comparison with a NAD-specific alcohol dehydrogenase (ADH-A) and a NADP-dependent AADH-B studied previously. Cultivated tetraploid emmer wheats (T. turgidum s. l.) and hexaploid bread wheats (T. aestivum s. l.) are all fixed for a heterozygous triplet, E^0.58/E^0.64. The slowest isoenzyme, E^0.58, is controlled by a homoeoallelic gene on the chromosome arm 6AL of T. aestivum cv. Chinese Spring and is inherent in all diploid wheats, T. monococcum s. Str., T. boeoticum s. l. and T. urartu. The fastest isoenzyme, E^0.64, is presumably controlled by the B- and D-genome homoeoalleles of the bread wheat and is the commonest alloenzyme of diploid goat-grasses, including Ae. speltaides and Ae. tauschii. The tetraploid T. timopheevii s. str. has a particular heterozygous triplet E^0.56/E^0.71, whereas the hexaploid T. zhukovskyi exhibited polymorphism with electromorphs characteristic of T. timopheevii and T. monococcum. Wild tetraploid wheats, T. dicoccoides and T. araraticum, showed partially homologous intraspecific variation of AADH-E with heterozygous triplets E^0.58/E^0.64 (the commonest), E^0.58/E^0.71, E^0.45/E^0.58, E^0.48/E^0.58 and E^0.56/E^0.58 recorded. Polyploid goatgrasses of the D-genome group, excepting Ae. cylindrica, are fixed for the common triplet E^0.58/E^0.64. Ae. cylindrica and polyploid goatgrasses of the C^u-genome group, excepting Ae. kotschyi, are homozygous for E^0.64. Ae. kotschyi is exceptional, showing fixed heterozygosity for both AADH-E and ADH-A with unique triplets E^0.56/E^0.64 and A^0.49/A^0.56.     

Marker-assisted development and characterization of a set of Triticum aestivum lines carrying different introgressions from the T. timopheevii genome

A set of common wheat introgression lines carrying one or two introgressions from Triticum timopheevii was produced by means of marker-assisted backcross selection. The starting material consisted of two BC1F20 (T. aestivum*2/T. timopheevii) lines with resistance to leaf rust, stem rust, powdery mildew, spot blotch, and loose smut and containing multiple 1A^t, 2A^t, 2G, 3A^tL, 3GL, 4GL, 5A^tL, 5GL, and 6G T. timopheevii chromosome fragments. The two lines were crossed with, and backcrossed three times to common wheat cultivar Saratovskaya 29. In total, 275 BC2F1 and BC3F2 plants were characterized by microsatellite markers and in situ hybridization. Molecular and cytological analyses revealed 38 plants with a single introgression from chromosomes 2G, 5GL, or 6G of T. timopheevii and 72 plants, each with two introgressions, among them three plants carrying a T. timopheevii translocation involving the D genome (2DS.2GL). It was observed that the lengths of fragments introgressed from the A^t genome were more than halved in the BC2 generation, while the lengths of 2G and 5GL introgressed fragments were only slightly reduced after the third backcross. The introgression lines were tested for resistance to the native Puccinia triticina population of the Western Siberian region of Russia. Lines with a single introgressed 5GL region carrying the major leaf rust resistance locus, QLr.icg-5B, were completely resistant. The presence of two minor resistance loci, QLr.icg-2A and QLr.icg-1A, suppressed disease development and reduced the number of urediniospores by up to 25 % but did not lead to a hypersensitive response. The introgression lines therefore constitute promising sources of new resistance to Puccinia triticina.     

Cytogenetic Analysis of Hybrids Resistant to Yellow Rust and Powdery Mildew Obtained by Crossing Common Wheat (Triticum aestivum L., AABBDD) with Wheats of the Timopheevi Group (AtAtGG)

The karyotypes of 47 hybrid lines obtained from crosses of common wheat Triticum aestivum L. (cv. Rodina and line 353) with Triticum timopheevii(Zhuk.) Zhuk. (A ^t A ^t GG) and related species T. militinae Zhuk. et Migusch. (A ^t A ^t GG) and T. kiharae Dorof. et Migusch. (A ^t A ^t GGD ^sq D ^sq) were analyzed by C-banding. Most lines were resistant to yellow rust and powdery mildew. The introgression of alien genetic material to the common wheat genome was realized via substitutions of complete A ⁺-,G-, and D-genome chromosomes, chromosome arms, or their fragments. The pattern of chromosome substitutions in resistant lines differed from that in introgressive hybrids selected for other traits. Substitutions of chromosomes 6G, 2A^t, 2G, and 5G were revealed in 31, 23, 18, and 13 lines, respectively. Substitutions of chromosomes 4G, 4A^t, and 6A^t were not observed. In 15 lines, a 5BS.5BL-5GL translocation was identified. High frequency of substitutions of chromosomes 2A^t, 2G, 5G, and 6G indicate that they may carry the resistance genes and that they are closely related to the respective homoeologous chromosomes of common wheat that determines their high compensation ability.     

Molecular cytogenetic characterization of <Emphasis Type="Italic">Triticum timopheevii</Emphasis> chromosomes provides new insight on genome evolution of <Emphasis Type="Italic">T. zhukovskyi</Emphasis>

Triticum timopheevii (2n = 4x = 28, GGA^tA^t) is a tetraploid wheat formerly cultivated in western Georgia. The natural allopolyploid Triticum zhukovskyi is a hexaploid taxon originated from hybridization of T. timopheevii with cultivated einkorn T. monococcum (2n = 2x = 14, A^mA^m). Karyotypically T. timopheevii and T. zhukovskyi differ from other tetraploid and hexaploid wheats and were assigned to the section Timopheevii of the genus Triticum L. Triticum timopheevii and T. zhukovskyi are resistant to many fungal diseases and therefore could potentially be utilized for wheat improvement. We were aiming to precisely identify all T. timopheevii chromosomes and to trace the evolution of T. zhukovskyi. For this, we developed a set of molecular cytogenetic landmarks based on eleven DNA probes. Each chromosome can now be characterized by two to eight probes. The pTa-535 sequence allows the identification of all A^t-genome chromosomes, whereas G-genome and some A^t-genome chromosomes can be identified using (GAA/CTT) _n and pSc119.2 probes. The probes pAesp_SAT86, pAs1, Spelt-1, Spelt-52 and 5S and 45S rDNA can be applied as additional markers to discriminate particular chromosomes or chromosomal regions. The distribution of (GAA/CTT) _n , pTa-535 and pSc119.2 DNA probes on T. timopheevii chromosomes is distinct from other tetraploid wheats and can therefore be used to track individual chromosomes in introgression programs. Our study confirms the origin of T. zhukovskyi from hybridization of T. timopheevii with T. monococcum; however, we show that the emergence was accompanied by changes involving mostly A^t-genome chromosomes. This may be due to the presence of two closely related A-genomes in the T. zhukovskyi karyotype.     

Repeated DNA sequences inTriticum (Poaceae): Chromosomal mapping and its bearing on the evolution of B and G genomes

The somatic chromosomes ofTriticum turgicum var.durum cv. Langdon andT. dicoccoides (AABB tetraploids),T. timopheevii, andT. araraticum (AAGG tetraploids) were assayed for distribution patterns of a highly repeated 120bp DNA sequence by in situ hybridization. The repeated sequence appears to be an ancient sequence shared withSecale andAegilops. The distribution patterns of the chromosomes were compared to the patterns of the A and B genome chromosomes ofT. aestivum cv. Chinese Spring (AABBDD hexaploid).T. turgidum andT. dicoccoides were observed to have identical in situ hybridization patterns. In both species, nine chromosomes with a total of 21 sites of hybridization were observed. The pattern, with few exceptions, was identical to that of Chinese Spring.T. araraticum andT. timopheevii were observed to have different patterns. InT. araraticum, six chromosomes with 21 total hybridization sites are present while inT. timopheevii nine chromosomes with 19 total sites exist. Major differences in hybridization patterns were observed between the B and G genomes. The divergence of the tetraploid wheats in this study appears to have resulted in changes in location, not in amount, of the ancient repeated sequence.     

Intergenomic chromosome substitutions in wheat interspecific hybrids and their use in the development of a genetic nomenclature of <Emphasis Type="Italic">Triticum timopheevii</Emphasis> chromosomes

The results of analysis of the genome formation in interspecific hybrids of Triticum aestivum with T. timopheevii are reviewed. The spectra of substitutions and rearrangements are shown to depend on the genotypes of the parental forms and on the direction of selection. The frequencies of substitutions of individual T. timopheevii chromosomes significantly vary and reflect the level of their divergence relative to the common wheat chromosomes. Some aspects of classification of the A^t- and G-genome chromosomes are discussed.     

Structural evolution of wheat chromosomes 4A, 5A,and 7B and its impact on recombination

The construction of comparative genetic maps of chromosomes 4A^m and 5A^m of Triticum monococcum and chromosomes of homoeologous groups 4, 5 and 7 of T. aestivum has provided insight into the evolution of these chromosomes. The structures of chromosomes 4A, 5A and 7B of modern-day hexaploid bread wheat can be explained by a 4AL/5AL translocation that occurred at the diploid level and is present both in T. monococcum and T. aestivum. Three further rearrangements, a 4AL/7BS translocation, a pericentric inversion and a paracentric inversion, have taken place in the tetraploid progenitor of hexaploid wheat. These structural rearrangements and the evolution of chromosomes 4A, 5A and 7B of bread wheat are discussed. The presence of the 4AL/5AL translocation in several Triticeae genomes raises two questions — which state is the more primitive, and is the translocation of mono- or poly-phylogenetic origin? The rearrangements that have occurred in chromosome 4A resulted in segments of both arms having different positions relative to the telomere, compared to 4A^m and to 4B and 4D. Comparisons of map length in these regions indicate that genetic length is a function of distance from the telomere, with the distal regions showing the highest recombination.     

Relationships betweenNor-loci from differentTriticeae species

TheNor-loci of polyploid wheats and their putative diploid progenitor species were assayed by probing isolated nuclear DNA with ribosomal DNA spacer sequences (spacer rDNA sequences, isolated by cloning), from theNor-loci of genomes B (Triticum aestivum), G (T. timopheevi), B (syn. S,T. speltoides), A (T. monococcum) and V (Dasypyrum villosum). DNA samples for analysis were digested with the restriction endonuclease Taq 1 and assayed by DNA-DNA hybridization under standard (37°C) and high stringency (64°C) conditions. The assay procedure emphasized differences between the divergent spacer sequences of the polyploid species and allowed relative homologies to the respective sequences in diploid species to be established. — The studies indicated thatT. timopheevi andT. speltoides contain different sets of spacer rDNA sequences which were readily distinguishable and, in the case ofT. timopheevi, assigned toNor-loci on different chromosomes. This contrast with the spacer rDNA sequences of the majorNor-loci on chromosomes 1 B and 6 B inT. aestivum, which were difficult to distinguish and were deduced to contain very similar sequences. Among the diploid progenitor species only the spacer rDNA fromT. speltoides shared close homology with polyploid wheat species. OneNor-locus inT. timopheevi (on chromosome 6 G) did not show close homology with any of the rDNA spacer probes available. — The data suggestsT. speltoides was the origin of someNor-loci for both theT. timopheevi andT. turgidum lines of tetraploid wheats. The possibility that the 6GNor-locus inT. timopheevi may have derived from an unknown diploid species by introgressive hybridization is discussed. The spacer rDNA sequence probe fromT. monococcum shared good homology with some accessions ofD. villosum and a line ofT. dicoccoides; the implications of this finding for evolution of present-day wheats are discussed.     

Molecular characterisation of the amino- and carboxyl-domains in different Glu-A1x alleles of Triticum urartu Thum. ex Gandil.

The wild diploid wheat (Triticum urartu Thum. ex Gandil.) is a potential gene source for wheat breeding, as this species has been identified as the A-genome donor in polyploid wheats. One important wheat breeding trait is bread-making quality, which is associated in bread wheat (T. aestivum ssp. aestivum L. em. Thell.) with the high-molecular-weight glutenin subunits. In T. urartu, these proteins are encoded by the Glu-A1x and Glu-A1Ay genes at the Glu-A ^u 1 locus. The Glu-A1x genes of 12 Glu-A ^u 1 allelic variants previously detected in this species were analysed using PCR amplification and sequencing. Data showed wide diversity for the Glu-A1x alleles in T. urartu, which also showed clear differences to the bread wheat alleles. This variation could enlarge the high-quality genetic pool of modern wheat and be used to diversify the bread-making quality in durum (T. turgidum ssp. durum Desf. em. Husn.) and common wheat.     

Genetic variability of the wild diploid wheat Triticum urartu revealed by RFLP and RAPD markers

 Genetic variability among 49 accessions of Triticum urartu was estimated by RFLP and RAPD marker analyses, and the two data sets were compared. One T. timopheevii accession and two accessions of T. durum and T. aestivum, respectively, were included to identify T. urartu accessions closely related to these polyploid wheats. Twenty eight RFLP clones and 29 RAPD primers generated 451 and 155 polymorphic bands, respectively. The three accessions from Armenia clustered together and were well separated from all other accessions, which showed less pronounced geographical patterns. Genetic similarity and co-phenetic values calculated with RAPD markers were very similar to those calculated with RFLP markers for the intraspecific comparisons, but not for the interspecific comparisons. The identification of individual T. urartu accessions which are more related to polyploid wheats than others was not possible. Received: 14 May 1996 / Accepted: 13 September 1996