Rapid Elimination of Low-Copy DNA Sequences in Polyploid Wheat: A Possible Mechanism for Differentiation of Homoeologous Chromosomes

To study genome evolution in allopolyploid plants, we analyzed polyploid wheats and their diploid progenitors for the occurrence of 16 low-copy chromosome- or genome-specific sequences isolated from hexaploid wheat. Based on their occurrence in the diploid species, we classified the sequences into two groups: group I, found in only one of the three diploid progenitors of hexaploid wheat, and group II, found in all three diploid progenitors. The absence of group II sequences from one genome of tetraploid wheat and from two genomes of hexaploid wheat indicates their specific elimination from these genomes at the polyploid level. Analysis of a newly synthesized amphiploid, having a genomic constitution analogous to that of hexaploid wheat, revealed a pattern of sequence elimination similar to the one found in hexaploid wheat. Apparently, speciation through allopolyploidy is accompanied by a rapid, nonrandom elimination of specific, low-copy, probably noncoding DNA sequences at the early stages of allopolyploidization, resulting in further divergence of homoeologous chromosomes (partially homologous chromosomes of different genomes carrying the same order of gene loci). We suggest that such genomic changes may provide the physical basis for the diploid-like meiotic behavior of polyploid wheat.     

An improved method of genomic in situ hybridization (GISH) for distinguishing closely related genomes of tetraploid and hexaploid wheat species

An improved modification of genomic in situ hybridization (GISH) was proposed. It allows clear and reproducible discrimination between closely related genomes of both tetraploid and hexaploid wheat species due to preannealing of labeled DNA probes and prehybridization of chromosomal samples with blocking DNA. The method was applied to analyze intergenomic translocations 6A:6B and 1A:6B identified in the IG46147 and IG116188 samples of tetraploid wheat Triticum dicoccoides by C-banding. The structure of the rearranged chromosomes was defined for two translocation variants, and the breakpoints were identified on the chromosome arms. Possible application of the developed GISH variant to study genome reorganizations during speciation of allopolyploid plants in evolution is discussed.     

Electrophoretic survey of seedling esterases in wheats in relation to their phylogeny

Summary Evolutionary and ontogenetic variation of six seedling esterases of independent genetic control is studied in polyploid wheats and their diploid relatives by means of polyacrylamide gel electrophoresis. Four of them are shown to be controlled by homoeoallelic genes in chromosomes of third, sixth and seventh homoeologous groups.The isoesterase electrophoretic data are considered supporting a monophyletic origin of both the primitive tetraploid and the primitive hexaploid wheat from which contemporary taxa of polyploid wheats have emerged polyphyletically and polytopically through recurrent introgressive hybridization and accumulation of mutations. Ancestral diploids belonging or closely related to Triticum boeoticum, T. urartu, Aegilops speltoides and Ae. tauschii ssp. strangulata are genetically the most suitable genome donors of polyploid wheats. Diploids of the Emarginata subsection of the section Sitopsis, Aegilops longissima s.str., Ae. sharonensis, Ae. searsii and Ae. bicornis, are unsuitable for the role of the wheat B genome donors, being all fixed for the esterase B and D electromorphs different from those of tetraploid wheats.     

普通小麦7B染色体两类低拷贝专化DNA序列的特性

研究表明 ,多倍体小麦基因组中存在一类低拷贝、染色体专化的DNA序列 ,其在多倍体形成时常表现出不稳定性。这类序列被认为在异源多倍体的建立和稳定中起着关键作用。为进一步研究这一问题 ,对通过染色体显微切割从普通小麦 (TriticumaestivumL .)中分离的 5个 7B染色体专化DNA序列的特性进行了研究。以这些序列为探针对大量的多倍体小麦和它们的二倍体祖先物种进行了Southern杂交分析。结果表明 ,这些序列可被分为两种类型 :其中的 4个序列与所有的多倍体物种均杂交 ,但是在二倍体水平上 ,它们却只与和多倍体小麦B基因组紧密相关的物种杂交 ,这说明这些序列是在二倍体物种分化以后产生的 ,然后垂直传递给多倍体 ;其中的 1个序列与所有的二倍体及多倍体物种均杂交 ,暗示在多倍体形成后这些序列从A和D基因组中消除了。用这一序列分别与一个人工合成的六倍体和四倍体小麦进行Southern杂交的结果表明 ,序列消除是一个迅速的事件而且很可能与这些序列的甲基化状态有关。认为这些低拷贝的染色体专化序列对于多倍体形成后部分同源染色体之间的进一步分化起着重要作用。     

Phylogenetic relationships of Triticum and Aegilops and evidence for the origin of the A, B, and D genomes of common wheat (Triticum aestivum)

Common wheat (Triticum aestivum) has for decades been a textbook example of the evolution of a major crop species by allopolyploidization. Using a sophisticated extension of the PCR technique, we have successfully isolated two single-copy nuclear genes, DMC1 and EF-G, from each of the three genomes found in hexaploid wheat (BA(u)D) and from the two genomes of the tetraploid progenitor Triticum turgidum (BA(u)). By subjecting these sequences to phylogenetic analysis together with sequences from representatives of all the diploid Triticeae genera we are able for the first time to provide simultaneous and strongly supported evidence for the D genome being derived from Aegilops tauschii, the A(u) genome being derived from Triticum urartu, and the hitherto enigmatic B genome being derived from Aegilops speltoides. Previous problems of identifying the B genome donor may be associated with a higher diversification rate of the B genome compared to the A(u) genome in the polyploid wheats. The phylogenetic hypothesis further suggests that neither Triticum, Aegilops, nor Triticum plus Aegilops are monophyletic.     

Expression of high zinc efficiency of Aegilops tauschii and Triticum monococcum in synthetic hexaploid wheats

The effect of varied zinc (Zn) supply on shoot and root dry matter production, severity of Zn deficiency symptoms and Zn tissue concentrations was studied in two Triticum turgidum (BBAA) genotypes and three synthetic hexaploid wheat genotypes by growing plants in a Zn-deficient calcareous soil under greenhouse conditions with (+Zn=5 mg kg^-1 soil) and without (−Zn) Zn supply. Two synthetic wheats (BBAADD) were derived from two different Aegilops tauschii (DD) accessions using same Triticum turgidum (BBAA), while one synthetic wheat (BBAAAA) was derived from Triticum turgidum (BBAA) and Triticum monococcum (AA). Visible symptoms of Zn deficiency, such as occurrence of necrotic patches on leaves and reduction in shoot elongation developed more rapidly and severely in tetraploid wheats than in synthetic hexaploid wheats. Correspondingly, decreases in shoot and root dry matter production due to Zn deficiency were higher in tetraploid wheats than in synthetic hexaploid wheats. Transfer of the DD genome from Aegilops tauschii or the AA genome from Triticum monococcum to tetraploid wheat greatly improved root and particularly shoot growth under Zn-deficient, but not under Zn-sufficient conditions. Better growth and lesser Zn deficiency symptoms in synthetic hexaploid wheats than in tetraploid wheats were not accompanied by increases in Zn concentration per unit dry weight, but related more to the total amount of Zn per shoot, especially in the case of synthetic wheats derived from Aegilops tauschii. This result indicates higher Zn uptake capacity of synthetic wheats. The results demonstrated that the genes for high Zn efficiency from Aegilops tauschii (DD) and Triticum monococcum (AA) are expressed in the synthetic hexaploid wheats. These wheat relatives can be used as valuable sources of genes for improvement of Zn efficiency in wheat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Recurrent deletions of puroindoline genes at the grain hardness locus in four independent lineages of polyploid wheat

Polyploidy is known to induce numerous genetic and epigenetic changes but little is known about their physiological bases. In wheat, grain texture is mainly determined by the Hardness (Ha) locus consisting of genes Puroindoline a (Pina) and b (Pinb). These genes are conserved in diploid progenitors but were deleted from the A and B genomes of tetraploid Triticum turgidum (AB). We now report the recurrent deletions of Pina-Pinb in other lineages of polyploid wheat. We analyzed the Ha haplotype structure in 90 diploid and 300 polyploid accessions of Triticum and Aegilops spp. Pin genes were conserved in all diploid species and deletion haplotypes were detected in all polyploid Triticum and most of the polyploid Aegilops spp. Two Pina-Pinb deletion haplotypes were found in hexaploid wheat (Triticum aestivum; ABD). Pina and Pinb were eliminated from the G genome, but maintained in the A genome of tetraploid Triticum timopheevii (AG). Subsequently, Pina and Pinb were deleted from the A genome but retained in the A(m) genome of hexaploid Triticum zhukovskyi (A(m)AG). Comparison of deletion breakpoints demonstrated that the Pina-Pinb deletion occurred independently and recurrently in the four polyploid wheat species. The implications of Pina-Pinb deletions for polyploid-driven evolution of gene and genome and its possible physiological significance are discussed.     

The synaptic behaviour of the wild forms of Triticum turgidum and T. timopheevii.

Different wild allopolyploid species of Triticeae show extensive bivalent formation at zygotene while a considerable number of multivalents is present in cultivated polyploid wheats. To study the chromosome behaviour at early meiotic stages in wild forms of tetraploid wheats Triticum turgidum and T timopheevii (2n = 4x = 28) we have analysed the synaptic pattern in fully traced spread nuclei at mid- and late zygotene and at pachytene of wild accessions of these species. The mean number of synaptonemal complex (SC) bivalents at mid-zygotene ranged from 12.22 to 13.14 among the accessions studied indicating a strong restriction of synapsis initiation to homologous chromosomes. The mean of bivalents increased at pachytene because of the transformation of multivalents into bivalents. Ring bivalents observed at metaphase I support that SC bivalents were formed by homologous chromosomes. The average values of SC bivalents at mid-zygotene in the wild forms are much higher than the average values observed in the cultivated tetraploid wheats but similar to that of a mutant line of T turgidum with a duplication that includes Ph1, the major homoeologous pairing suppressor locus. These results suggest that the efficiency of the mechanism operating in the homologous recognition for synapsis is higher in wild wheat populations than in cultivated varieties. Apparently, a relatively detrimental modification of the pairing regulating genetic system accompanied the domestication of the wild wheat forms.     

Quantification of genetic relationships among A genomes of wheats.

The genetic relationships of A genomes of Triticum urartu (Au) and Triticum monococcum (Am) in polyploid wheats are explored and quantified by AFLP fingerprinting. Forty-one accessions of A-genome diploid wheats, 3 of AG-genome wheats, 19 of AB-genome wheats, 15 of ABD-genome wheats, and 1 of the D-genome donor Ae. tauschii have been analysed. Based on 7 AFLP primer combinations, 423 bands were identified as potentially A genome specific. The bands were reduced to 239 by eliminating those present in autoradiograms of Ae. tauschii, bands interpreted as common to all wheat genomes. Neighbour-joining analysis separates T. urartu from T. monococcum. Triticum urartu has the closest relationship to polyploid wheats. Triticum turgidum subsp. dicoccum and T. turgidum subsp. durum lines are included in tightly linked clusters. The hexaploid spelts occupy positions in the phylogenetic tree intermediate between bread wheats and T. turgidum. The AG-genome accessions cluster in a position quite distant from both diploid and other polyploid wheats. The estimates of similarity between A genomes of diploid and polyploid wheats indicate that, compared with Am, Au has around 20% higher similarity to the genomes of polyploid wheats. Triticum timo pheevii AG genome is molecularly equidistant from those of Au and Am wheats.     

Meiotic behaviour of tetraploid wheats (<Emphasis Type="Italic">Triticum turgidum</Emphasis> L.) and their synthetic hexaploid wheat derivates influenced by meiotic restitution and heat stress

Meiotic restitution is considered to be a common mechanism of polyploidization in plants and hence is one of the most important processes in plant speciation. Meiotic behaviour of plant chromosomes is influenced by both genetic and environmental factors. In this study, the meiotic behaviour of cereal crops was investigated, which includes tetraploid wheat genotypes (with and without the meiotic restitution trait) and their derivates (synthetic hexaploid wheats and a doubled haploid (DH) line), grown at two planting dates in the field. In addition, two local landraces of emmer wheat (Triticum turgidum ssp. dicoccum), one wheat cultivar (Chinese spring), one DH triticale cultivar (Eleanor) and one rye accession were included. Immature spikes of mid-autumn and end-winter sowing plants were collected in April and May 2008, respectively, fixed in Carnoy’s solution and stained with hematoxylin. Pollen mother cells (PMCs) from anthers at different stages of meiotic process were analysed for their chromosomal behaviour and irregularities. Meiotic aberrations such as laggards, chromosome bridges, micronuclei, abnormal cytokines, chromatin pulling and meiotic restitution were observed and the studied genotypes were accordingly ranked as follows: triticale > synthetic hexaploid wheats > tetraploid wheats possessing meiotic restitution > tetraploid wheats lacking meiotic restitution > rye. The results indicated that the samples that had been planted in the autumn, thus experiencing an optimum temperature level at the flowering stage, exhibited less meiotic irregularities than winter planting samples that encountered heat stress at the flowering period.     

Molecular phylogeny of the genus Triticum L

The genus Triticum L. includes the major cereal crop, common or bread wheat (hexaploid Triticum aestivum L.), and other important cultivated species. Here, we conducted a phylogenetic analysis of all known wheat species and the closely related Aegilops species. This analysis was based on chloroplast matK gene comparison along with trnL intron sequences of some species. Polyploid wheat species are successfully divided only into two groups – Emmer (sections Dicoccoides and Triticum) and Timopheevii (section Timopheevii). Results reveal strictly maternal plastid inheritance of synthetic wheat amphiploids included in the study. A concordance of chloroplast origin with the definite nuclear genomes of polyploid species that were inherited at the last hybridization events was found. Our analysis suggests that there were two ancestral representatives of Aegilops speltoides Tausch that participated in the speciation of polyploid wheats with B and G genome in their genome composition. However, G genome species are younger in evolution than ones with B genome. B genome-specific PCR primers were developed for amplification of Acc-1 gene.     

Phytosiderophore release in Aegilops tauschii and Triticum species under zinc and iron deficiencies.

Using three diploid (Triticum monococcum, AA), three tetraploid (Triticum turgidum, BBAA), two hexaploid (Triticum aestivum and Triticum compactum, BBAADD) wheats and two Aegilops tauschii (DD) genotypes, experiments were carried out under controlled environmental conditions in nutrient solution (i) to study the relationships between the rates of phytosiderophore (PS) release from the roots and the tolerance of diploid, tetraploid, and hexaploid wheats and AE: tauschii to zinc (Zn) and iron (Fe) deficiencies, and (ii) to assess the role of different genomes in PS release from roots under different regimes of Zn and Fe supply. Phytosiderophores released from roots were determined both by measurement of Cu mobilized from a Cu-loaded resin and identification by using HPLC analysis. Compared to tetraploid wheats, diploid and hexaploid wheats were less affected by Zn deficiency as judged from the severity of leaf symptoms. Aegilops tauschii showed very slight Zn deficiency symptoms possibly due to its slower growth rate. Under Fe-deficient conditions, all wheat genotypes used were similarly chlorotic; however, development of chlorosis was first observed in tetraploid wheats. Correlation between PS release rate determined by Cu-mobilization test and HPLC analysis was highly significant. According to HPLC analysis, all genotypes of Triticum and AE: tauschii species released only one PS, 2'-deoxymugineic acid, both under Fe and Zn deficiency. Under Zn deficiency, rates of PS release in tetraploid wheats averaged 1 micromol x (30 plants)(-1) x (3 h)(-1), while in hexaploid wheats rate of PS release was around 14 micromol x (30 plants)(-1) x (3 h)(-1). Diploid wheats and AE: tauschii accessions behaved similarly in their capacity to release PS and intermediate between tetraploid and hexaploid wheats regarding the PS release capacity. All Triticum and Aegilops species released more PS under Fe than Zn deficiency, particularly when the rate of PS release was expressed per unit dry weight of roots. On average, the rates of PS release under Fe deficiency were 3.0, 5.7, 8.4, and 16 micromol x (30 plants)(-1) x (3 h)(-1) for AE: tauschii, diploid, tetraploid and hexaploid wheats, respectively. The results of the present study show that the PS release mechanism in wheat is expressed effectively when three genomes, A, B and D, come together, indicating complementary action of the corresponding genes from A, B and D genomes to activate biosynthesis and release of PS.     

The presence of the enhanced K/Na discrimination trait in diploid Triticum species

Summary A number of accessions of the three species of diploid wheat, Triticum boeoticum, T. monococcum, and T. urartu, were grown in 50 mol m^-3 NaCl+2.5 mol m^-3 CaCl₂. Sodium accumulation in the leaves was low and potassium concentrations remained high. This was not the case in T. durum grown under the same conditions, and indicates the presence in diploid wheats of the enhanced K/Na discrimination character which has previously been found in Aegilops squarrosa and hexaploid wheat. None of the accessions of diploid wheat showed poor K/Na discrimination, which suggests that if the A genome of modern tetraploid wheats was derived from a diploid Triticum species, then the enhanced K/Na discrimination character became altered after the formation of the original allopolyploid. Another possibility is that a diploid wheat that did not have the enhanced K/Na discrimination character was involved in the hybridization event which produced tetraploid wheat, and that this diploid is now extinct or has not yet been discovered.     

Molecular diversity at 18 loci in 321 wild and 92 domesticate lines reveal no reduction of nucleotide diversity during Triticum monococcum (Einkorn) domestication: implications for the origin of agriculture

The diploid wheat Triticum monococcum L. (einkorn) was among the first crops domesticated by humans in the Fertile Crescent 10,000 years ago. During the last 5,000 years, it was replaced by tetraploid and hexaploid wheats and largely forgotten by modern breeders. Einkorn germplasm is thus devoid of breeding bottlenecks and has therefore preserved in unfiltered form the full spectrum of genetic variation that was present during its domestication. We investigated haplotype variation among >12 million nucleotides sequenced at 18 loci across 321 wild and 92 domesticate T. monococcum lines. In contrast to previous studies of cereal domestication, we sampled hundreds of wild lines, rather than a few dozen. Unexpectedly, our broad sample of wild lines reveals that wild einkorn underwent a process of natural genetic differentiation, most likely an incipient speciation, prior to domestication. That natural differentiation was previously overlooked within wild einkorn, but it bears heavily upon inferences concerning the domestication process because it brought forth 3 genetically, and to some extent morphologically, distinct wild einkorn races that we designate here as alpha, beta, and gamma. Only one of those natural races, beta, was exploited by humans for domestication. Nucleotide diversity and haplotype diversity in domesticate einkorn is higher than in its wild sister group, the einkorn beta race, indicating that einkorn underwent no reduction of diversity during domestication. This is in contrast to findings from previous studies of domestication history among more intensely bred crop species. Taken together with archaeological findings from the Fertile Crescent, the data indicate that a specific wild einkorn race that arose without human intervention was subjected to multiple independent domestication events.     

小麦染色体组的起源与进化探讨

对小麦染色体组的起源及其进化进行了全面综述后,提出了一个新的小麦进化途径,并认为：（１）Ｔｒｉｔｉｃｕｍｍｏｎｏｃｏｃｕｍｖａｒｕｒａｒｔｕ是多倍体小麦Ａ组的原初供体,在Ａ组进入多倍体小麦后有Ｔｍｏｎｏｖａｒｂｏｅｏｔｉｃｕｍ的基因渗入;（２）Ｂ和Ｇ组的原初供体是Ｔｓｐｅｌｔｏｉｄｅｓ的Ｓ组,在该Ｓ组进入多倍体小麦后有两个进化方向,即Ｓ组结构分化形成Ｇ组和Ｓ组经外源染色体代换及重组等而进化成Ｂ组;（３）Ｔｔｕｒｇｉｄｕｍ和Ｔｔｉｍｏｐｈｅｖｉ都是来自Ｔｓｐｅｌｔｏｉｄｅｓ为母本与Ｔｍｏｎｏｖａｒｕｒａｒｔｕ杂交后并双二倍化而形成的原初四倍体小麦（ＳＳＡＡ）,并由它分别经遗传渗入和结构分化而成;（４）Ｔｚｈｕｋｏｖｓｋｙｉ是Ｔｔｉｍｏｐｈｅｖｉ作母本与Ｔｍｏｎｏｖａｒｂｏｅｏｔｉｃｕｍ杂交并双二倍化而形成,故它具有分别来自Ｔｍｏｎｏｖａｒｕｒａｒｔｕ和Ｔｍｏｎｏｖａｒｂｏｅｏｔｉｃｕｍ的两类Ａ组;（５）Ｔａｅｓｔｉｖｕｍ的Ｄ组来自Ｔｔａｕｓｃｈｉ;（６）无论Ａ组、Ｂ组、Ｄ组、Ｇ组在进入多倍体小麦后均有相当分化,同时在其供体种中也有一定分化。     

Electrophoretic analysis of substrate specificity of wheat alcohol dehydrogenases]

Electrophoresis in polyacrylamide gel slabs has been used to study the isoform composition and substrate specificity of alcohol dehydrogenases in the embryo and young seedlings of the diploid wheat Triticum monococcum L., the tetraploid T. dicoccon (Schrank) Schuebl and the hexaploid T. spelta L. Three alcohol dehydrogenases of different substrate specificity and developmental pattern were distinguished: a) the NAD-dependent alcohol dehydrogenase, catalyzing the oxidation of different primary and secondary aliphatic and aromatic alcohols, as well as certain compounds with several hydroxyl groups (tris, triethanolamin) and revealing, after electrophoresis, one major band in the diploid wheat and three bands in both polyploid wheats; b) the NADP-dependent aromatic alcohol dehydrogenase (substrate--cinnamic alcohol), revealing, after electrophoresis, one major fast moving band in the diploid wheat and two bands in polyploid wheats; c) an aromatic alcohol dehydrogenase (2-3 bands after electrophoreis) with no specificity to the cofactors (NAD or NADP).     

Differences in Zinc Efficiency among and within Diploid, Tetraploid and Hexaploid Wheats

Greenhouse experiments were carried out with six diploid, ninetetraploid and seven hexaploid wheats, including wild and primitivegenotypes, to study the influence of varied zinc (Zn) supplyon the severity of Zn deficiency symptoms, shoot dry matterproduction and shoot Zn concentrations. In addition to wildand primitive genotypes, one modern tetraploid cultivar withhigh sensitivity to Zn deficiency and two modern hexaploid cultivars,one highly sensitive to and one resistant to Zn deficiency,were included for comparison. Plants were grown for 44 d ina severely Zn-deficient calcareous soil, with (+Zn; 5 mg Znkg^-1soil) and without (-Zn) Zn fertilization. Visible Zn deficiencysymptoms, including whitish-brown necrotic patches on leaf blades,appeared very rapidly and severely in all tetraploid wheat genotypes.Compared with tetraploid wheats, diploid and hexaploid wheatswere less sensitive to Zn deficiency. With additional Zn, shootdry matter production was higher in tetraploid than diploidand hexaploid wheats. However, under Zn-deficient conditionstetraploid wheats had the lowest shoot dry matter production,indicating the very high sensitivity of tetraploid wheats toZn deficiency. Consequently, Zn efficiency expressed as theratio of shoot dry matter produced under Zn deficiency to Znfertilization, was much lower in tetraploid wheats than in diploidand hexaploid wheats. On average, Zn efficiency ratios were36% for tetraploid, 60% for diploid and 64% for hexaploid wheats.Differences in Zn efficiency among and within diploid, tetraploidand hexaploid wheats were positively related to the amount ofZn per shoot of the genotypes, but not to the amount of Zn perunit dry weight of shoots or seeds used in the experiments.The seeds of the accessions of tetraploid wild wheats containedup to 120 mg Zn kg^-1, but the resulting plants showed very highsensitivity to Zn deficiency. By contrast, hexaploid wheatsand primitive diploid wheats with much lower Zn concentrationsin seeds had higher Zn efficiencies. It is suggested that notonly enhanced Zn uptake capacity but also enhanced internalZn utilization capacity of genotypes play important roles indifferential expression of Zn efficiency. The results of thisstudy also suggest the importance of the A and D genomes asthe possible source of genes determining Zn efficiency in wheat.Copyright 1999 Annals of Botany Company Seeds, Triticum aestivum, Triticum monococcum, Triticum turgidum, zinc concentrations, zinc deficiency, zinc efficiency.     

Studies on the origin and evolution of tetraploid wheats based on the internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA

In this study, the internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA in the tetraploid wheats, Triticum turgidum (AABB) and Triticum timopheevii (AAGG), their possible diploid donors, i.e., Triticum monococcum (AA), Triticum urartu (AA), and five species in Aegilops sect. Sitopsis (SS genome), and a related species Aegilops tauschii were cloned and sequenced. ITS1 and ITS2 regions of 24 clones from the above species were compared. Phylogenetic analysis demonstrated that Aegilops speltoides was distinct from other species in Aegilops sect. Sitopsis and was the most-likely donor of the B and G genomes to tetraploid wheats. Two types of ITS repeats were cloned from Triticum turgidum ssp. dicoccoides, one markedly similar to that from T. monococcum ssp. boeoticum (AA), and the other to that from Ae. speltoides (SS). The former might have resulted from a recent integression event. The results also indicated that T. turgidum and T. timopheevii might have simultaneously originated from a common ancestral tetraploid species or be derived from two hybridization events but within a very short interval time. ITS paralogues in tetraploid wheats have not been uniformly homogenized by concerted evolution, and high heterogeneity has been found among repeats within individuals of tetraploid wheats. In some tetraploid wheats, the observed heterogeneity originated from the same genome (B or G). Three kinds of ITS repeats from the G genome of an individual of T. timopheevii ssp. araraticum were more divergent than that from inter-specific taxa. This study also demonstrated that hybridization and polyploidization might accelerate the evolution rate of ITS repeats in tetraploid wheats.     

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.     

Revisiting meiotic pairing in wheat synthetics in relation to the evolution of the meiotic system in wheat

A large panel of hexaploid wheat synthetics was developed. Their tetraploid parents consisted of either four extracted wheat tetraploids (ETWs) or four natural present-day tetraploids, and their diploid parents consisted of twenty accessions of Aegilops tauschii. Analysis of meiotic behaviour of the synthetics showed that chromosome pairing is highly variable and depends on the progenitor. The meiotic behaviour in the four ETWs was compared to that of the natural tetraploid wheats. It appears there was no evolution at the hexaploid level of the meiotic genes carried by the A and B genomes. We also reach the conclusion that the neo-allohexaploids at the origin of present-day wheat had a meiotic behaviour close to that of the present-day hexaploid wheat. It is likely that other neo-hexaploids with an impaired meiosis were formed, but they had no future due to their more or less rapid disappearance due to increasing aneuploidy level and structural changes, mainly Robertsonian translocations.