Distribution of Nonstructural Variation along Three Chromosome Arms between Wheat Cultivars Chinese Spring and Cheyenne

Metaphase I (MI) pairing of wheat homologous chromosomes is usually reduced in hybrids between cultivars relative to the parental inbred lines. Previous work suggested that this phenomenon is caused by polymorphism in nucleotide sequences (nonstructural chromosome variation) among wheat cultivars. The present work investigated the distribution of this variation along three selected chromosome arms between cultivars Chinese Spring and Cheyenne. Chinese Spring ditelosomics 3Aq, 6Ap and 6Bp were crossed with disomic substitutions of Cheyenne chromosomes 3A, 6A and 6B in Chinese Spring, respectively. The resulting F₁ plants, called substituted monotelodisomics, were crossed with the respective Chinese Spring monosomics, producing potentially "recombinant" substituted monosomics. When these "recombinant" chromosomes were combined with the parental Chinese Spring telosomes, marked reductions in mean telosome-pairing frequency were found compared with the corresponding Chinese Spring monotelodisomics. The mean pairing frequencies of the "recombinant" chromosomes showed a continuous distribution between those of the substituted and Chinese Spring monotelodisomics. The results suggest that the nonstructural variation that reduces MI pairing between chromosomes of different wheat cultivars is not localized in a specific site but distributed along each chromosome arm. Little variation was found among monotelodisomics for either the number of ring bivalents per cell or the number of univalents other than those constituting the heteromorphic pair. This implies that the reductions in MI pairing between the Cheyenne and Chinese Spring chromosomes are caused by something residing within these specific chromosomes that does not affect the pairing of the remaining Chinese Spring chromosomes in the same cell. Furthermore, the absence of parental types among the "recombinant"-substituted monotelodisomics suggests that the sequences involved in the variation studied here are capable of converting heterohomologous chromosomes to something intermediate in nature in the span of only a single generation.     

Distribution of Nonstructural Variation between Wheat Cultivars along Chromosome Arm 6Bp: Evidence from the Linkage Map and Physical Map of the Arm

Metaphase I (MI) pairing of homologous chromosomes in wheat intercultivar hybrids (heterohomologous chromosomes) is usually reduced relative to that within the inbred parental cultivars (euhomologous chromosomes). It was proposed elsewhere that this phenomenon is caused by polymorphism in nucleotide sequences (nonstructural chromosome variation) among wheat cultivars. The distribution of this polymorphism along chromosome arm 6Bp (=6BS) of cultivars Chinese Spring and Cheyenne was investigated. A population of potentially recombinant chromosomes derived from crossing over between telosome 6Bp of Chinese Spring and Cheyenne chromosome 6B was developed in the isogenic background of Chinese Spring. The approximate length of the Chinese Spring segment present in each of these chromosomes was assessed by determining for each chromosome the interval in which crossing over occurred (utilizing the rRNA gene region, a distal C-band and the gliadin gene region as markers). The MI pairing frequencies of these chromosomes (only the complete chromosomes were used) with the normal Chinese Spring telosome 6Bp were determined. These were directly proportional to the length of the euhomologous segment. The longer the incorporated euhomologous segment the better was the MI pairing. This provided evidence that the heterohomologous chromosomes are differentiated from each other in numerous sites distributed throughout the arm.—The comparison of the physical map of arm 6Bp with the linkage map showed a remarkable distortion of the linkage map; no crossing over was detected in the proximal 68% of the arm. A population of 49 recombinant chromosomes was assayed for recombination within the rRNA gene region, but none was detected. No new length variants of the nontranscribed spacer separating the 18S and 26S rRNA genes were detected either.     

Evidence for AEGILOPS SHARONENSIS Eig as the Donor of the B Genome of Wheat

A number of lines of evidence are advanced for the candidacy of Aegilops sharonensis Eig as the donor of the B genome of wheat. The cytoplasm of Ae. sharonensis is compatible with tetraploid wheat Triticum turgidum dicoccoides, as evidenced by the high level of chromosome pairing and fertility of the amphiploid Ae. sharonensis x T. turgidum dicoccoides. Ae. sharonensis chromosomes exhibit high levels of pairing with those of the B genome of wheat in hybrids with Ph-deficient hexaploid wheat and low levels of homoeologous pairing with T. monococcum chromosomes.——The amphidiploid between Ae. sharonensis and T. monococcum is very similar to T. turgidum dicoccoides in spike, spikelet and grain morphology. The karyotype of Ae. sharonensis resembles more closely that of extrapolated B genome karyotypes of wheat than do the karyotypes of other proposed B-genome donor species of Aegilops. Because of distinctiveness in cytological affinity and karyotype morphology between Ae. sharonensis and Ae. longissima, a separate genome symbol S^sh is proposed for the former species.     

Nonstructural Chromosome Differentiation among Wheat Cultivars, with Special Reference to Differentiation of Chromosomes in Related Species

Wheat cultivar Chinese Spring (Triticum aestivum L. em. Thell.) was crossed with cultivars Hope, Cheyenne and Timstein. In all three hybrids, the frequencies of pollen mother cells (PMCs) with univalents at metaphase I (MI) were higher than those in the parental cultivars. No multivalents were observed in the hybrids, indicating that the cultivars do not differ by translocations. Thirty-one Chinese Spring telosomic lines were then crossed with substitution lines in which single chromosomes of the three cultivars were substituted for their Chinese Spring homologues. The telosomic lines were also crossed with Chinese Spring. Data were collected on the frequencies (% of PMCs) of pairing of the telesomes with their homologues at MI and the regularity of pairing of the remaining 20 pairs of Chinese Spring chromosomes in the monotelodisomics obtained from these crosses. The reduced MI pairing in the intercultivar hybrids was caused primarily by chromosome differentiation, rather than by specific genes. Because the differentiation involved a large part of the chromosome complement in each hybrid, it was concluded that it could not be caused by structural changes such as inversions or translocations. In each case, the differentiation appeared to be unevenly distributed among the three wheat genomes. It is proposed that the same kind of differentiation, although of greater magnitude, differentiates homoeologous chromosomes and is responsible, together with structural differentiation, for poor chromosome pairing in interspecific hybrids.     

Homoeologous relationships of Aegilops speltoides chromosomes to bread wheat

 Homoeologous pairing at metaphase I was analysed in the standard-type, ph2b and ph1b hybrids of Triticum aestivum (AABBDD) and Aegilops speltoides (SS). Data from relative pairing affinities were used to predict homoeologous relationships of Ae. speltoides chromosomes to wheat. Chromosomes of both species, and their arms, were identified by C-banding. The Ae. speltoides genotype carried genes that induced a high level of homoeologous pairing in the three types of hybrids analyzed. All arms of the seven chromosomes of the S genome showed normal homoeologous pairing, which implies that no apparent chromosome rearrangements occurred in the evolution of Ae. speltoides relative to wheat. A pattern of preferential pairing of two types, A-D and B-S, confirmed that the S genome is very closely related to the B genome of wheat. Although this pairing pattern was also reported in hybrids of wheat with Ae. longissima and Ae. sharonensis, a different behaviour was found in group 5 chromosomes. In the hybrids of Ae. speltoides, chromosome 5B-5S pairing was much more frequent than 5D-5S, while these chromosome associations reached similar frequencies in the hybrids of Ae. longissima and Ae. sharonensis. These results are in agreement with the hypothesis that the B genome of wheat is derived from Ae. speltoides. Received: 8 January 1998 / Accepted: 4 February 1998     

Comparative genetic analysis of the Aegilops longissima and Ae. sharonensis genomes with common wheat

Aegilops longissima Schw. et Musch. (2n= 2x=14, S^lS^l) and Aegilops sharonensis Eig. (2n=2x=14, S^lS^l) are diploid species belonging to the section Sitopsis in the tribe Triticeae and potential donors of useful genes for wheat breeding. A comparative genetic map was constructed of the Ae. longissima genome, using RFLP probes with known location in wheat. A high degree of conserved colinearity was observed between the wild diploid and basic wheat genome, represented by the D genome of cultivated wheat. Chromosomes 1S^l, 2S^l, 3S^l, 5S^l and 6S^l are colinear with wheat chromosomes 1D, 2D, 3D, 5D and 6D, respectively. The analysis confirmed that chromosomes 4S^l and 7S^l are translocated relative to wheat. The short arms and major part of the long arms are homoeologous to most of wheat chromosomes 4D and 7D respectively, but the region corresponding to the distal segment of 7D was translocated from 7S^lL to the distal region of 4S^lL. The map and RFLP markers were then used to analyse the genomes and added chromosomes in a set of ’Chinese Spring’ (CS)/Ae. longissima chromosome additions. The study confirmed the availability of disomic CS/Ae. longissima addition lines for chromosomes 1S^l, 2S^l, 3S^l, 4S^l and 5S^l. An as yet unpublished set of Ae. sharonensis chromosome addition lines were also available for analysis. Due to the gametocidal nature of Ae. sharonensis chromosomes 2S^l and 4S^l, additions 1S^l, 3S^l, 5S^l, 6S^l and 7S^l were produced in a (4D)4S^l background, and 2S^l and 4S^l in a euploid wheat background. The analysis also confirmed that the 4/7 translocation found in Ae. longissima was not present in Ae. sharonensis although the two wild relatives of wheat are considered to be closely related. The phenotypes of the Ae. sharonensis addition lines are described in an Appendix. Received: 28 September 2000 / Accepted: 19 January 2001     

Cytogenetic and molecular mapping of the wheat-Aegilops longissima chromatin breakpoints in powdery mildew-resistant introgression lines

The amount of alien chromatin introgressed in eight wheat/Ae. longissima Pm13 recombinant lines, involving breakpoints on the short arms of wheat chromosomes 3B and 3D, was evaluated by cytogenetic and molecular approaches. For each line the residual homologous synaptic ability of the recombinant chromosome in its proximal wheat and distal alien portion was estimated through meiotic analyses. Subsequently, telocentric and RFLP mapping were used to assess the genetic distance from the wheat centromere to the wheat/Ae. longissima breakpoints. One 3B recombinant line was distinguished from the other four by the chromosome pairing and telocentric mapping analyses. RFLP analysis succeeded in differentiating the remaining four lines into two groups. Chromosome pairing and telocentric mapping of the three 3D recombinant lines suggested that all had distinct breakpoints. However, the RFLP data could not discriminate between the two more proximal translocations. Physical locations for some RFLP loci were determined by a comparison of genotypes and C-banding karyotypes. This showed a considerable expansion of the genetic map compared to its physical length.     

Development of a complete set of Triticum aestivum-Aegilops speltoides chromosome addition lines

Aegilops speltoides Tausch (2n = 2x = 14, SS) is considered as the closest living relative of the B and G genomes of polyploid wheats. A complete set of Triticum aestivum L. cv Chinese Spring-Ae. speltoides whole chromosomes and seven telosomic addition lines was established. A low pairing accession was selected for the isolation of the chromosome addition lines. Except for chromosomes 3S and 6S, which are presently only available as monosomic additions, all other lines were recovered as disomic or ditelosomic additions. The individual Ae. speltoides chromosomes isolated in the wheat background were assayed for their genetic effects on plant phenotype and cytologically characterized in terms of chromosome length, arm ratio, distribution of marker C-bands, and FISH sites using a Ae. speltoides-specific repetitive element, Gc1R-1, as a probe. The homoeology of the added Ae. speltoides chromosomes was established by using a standard set of RFLP probes. No chromosomal rearrangements relative to wheat were detected. Received: 28 June 1999 / Accepted: 16 November 1999     

Control of Endosperm Proteins in TRITICUM AESTIVUM (Var. Chinese Spring) and AEGILOPS UMBELLULATA by Homoeologous Group 1 Chromosomes

The genetic control of major wheat endosperm proteins by homoeologous group 1 chromosomes has been studied by two-dimensional polyacrylamide gel electrophoresis. The control of at least 15 distinct protein subunits or groups of protein subunits has been allocated to chromosomes 1A, 1B and 1D of Chinese Spring wheat from the analysis of grains of aneuploid genotypes. In addition, six protein subunits have been shown to be controlled by chromosome 1C^u of the related species, Aegilops umbellulata, from studies of wheat lines carrying disomic substitutions of 1C^u chromosomes. On the basis of protein subunit patterns, chromosome 1C^u is more closely related to chromosome 1D of wheat than to chromosomes 1A or 1B.     

Chromosome and nucleotide sequence differentiation in genomes of polyploid Triticum species

Summary The nature of genome change during polyploid evolution was studied by analysing selected species within the tribe Triticeae. The levels of genome changes examined included structural alterations (translocations, inversions), heterochromatinization, and nucleotide sequence change in the rDNA regions. These analyses provided data for evaluating models of genome evolution in polyploids in the genus Triticum, postulated on the basis of chromosome pairing at metaphase I in interspecies hybrids.The significance of structural chromosome alterations with respect to reduced MI chromosome pairing in interspecific hybrids was assayed by determining the incidence of heterozygosity for translocations and paracentric inversions in the A and B genomes of T. timopheevii ssp. araraticum (referred to as T. araraticum) represented by two lines, 1760 and 2541, and T. aestivum cv. Chinese Spring. Line 1760 differed from Chinese Spring by translocations in chromosomes 1A, 3A, 4A, 6A, 7A, 3B, 4B, 7B and possibly 2B. Line 2541 differed from Chinese Spring by translocations in chromosomes 3A, 6A, 6B and possibly 2B. Line 1760 also differed from Chinese Spring by paracentric inversions in arms 1AL and 4AL whereas line 2541 differed by inversions in 1BL and 4AL (not all chromosomes arms were assayed). The incidence of structural changes in the A and B genomes did not coincide with the more extensive differentiation of the B genomes relative to the A genomes as reflected by chromosome pairing studies.To assay changing degrees of heterochromatinization among species of the genus Triticum, all the diploid and polyploid species were C-banded. No general agreement was observed between the amount of heterochromatin and the ability of the respective chromosomes to pair with chromosomes of the ancestral species. Marked changes in the amount of heterochromatin were found to have occurred during the evolution of some of the polyploids.The analysis of the rDNA region provided evidence for rapid fixation of new repeated sequences at two levels, namely, among the 130 bp repeated sequences of the spacer and at the level of the repeated arrays of the 9 kb rDNA units. These occurred both within a given rDNA region and between rDNA regions on nonhomologous chromosomes. The levels of change in the rDNA regions provided good precedent for expecting extensive nucleotide sequence changes associated with differentiation of Triticum genomes and these processes are argued to be the principal cause of genome differentiation as revealed by chromosome pairing studies.     

Syntenic Relationships between the U and M Genomes of Aegilops,Wheat and the Model Species Brachypodium and Rice as Revealed by COS Markers

Diploid Aegilops umbellulata and Ae. comosa and their natural allotetraploid hybrids Ae. biuncialis and Ae. geniculata are important wild gene sources for wheat. With the aim of assisting in alien gene transfer, this study provides gene-based conserved orthologous set (COS) markers for the U and M genome chromosomes. Out of the 140 markers tested on a series of wheat-Aegilops chromosome introgression lines and flow-sorted subgenomic chromosome fractions, 100 were assigned to Aegilops chromosomes and six and seven duplications were identified in the U and M genomes, respectively. The marker-specific EST sequences were BLAST-ed to Brachypodium and rice genomic sequences to investigate macrosyntenic relationships between the U and M genomes of Aegilops, wheat and the model species. Five syntenic regions of Brachypodium identified genome rearrangements differentiating the U genome from the M genome and from the D genome of wheat. All of them seem to have evolved at the diploid level and to have been modified differentially in the polyploid species Ae. biuncialis and Ae. geniculata. A certain level of wheat–Aegilops homology was detected for group 1, 2, 3 and 5 chromosomes, while a clearly rearranged structure was showed for the group 4, 6 and 7 Aegilops chromosomes relative to wheat. The conserved orthologous set markers assigned to Aegilops chromosomes promise to accelerate gene introgression by facilitating the identification of alien chromatin. The syntenic relationships between the Aegilops species, wheat and model species will facilitate the targeted development of new markers specific for U and M genomic regions and will contribute to the understanding of molecular processes related to allopolyploidization.     

Investigation of Homologous Crossing over and Sister Chromatid Exchange in the Wheat Nor-B2 Locus Coding for Rrna and Gli-B2 Locus Coding for Gliadins

Recombination was investigated within the Nor-B2 locus of wheat chromosome 6B that contains several thousand of the 18S-5.8S-26S rRNA (rDNA) repeated units. Additionally, recombination was assessed for several chromosome regions, in arm 6Bq between the centromere and the B2 locus (awn suppressor) and in arm 6Bp between the centromere and Nor-B2, between Nor-B2 and a distal C-band and between Nor-B2 and Gli-B2 coding for gliadins. The experimental design permitted the distinction between crossing over between homologous chromosomes and exchange between sister chromatids. No homologous crossing over within the Nor-B2 locus was found in a sample of 446 chromosomes, but one exchange with the attributes of unequal sister chromatid exchange was identified. The molecular characteristics of this presumed sister chromatid exchange indicate that the spacer variants present in the Nor-B2 locus are clustered. No homologous recombination was detected within the distal Gli-B2 locus containing repeated genes coding for gliadin seed-storage proteins. Both arms of chromosome 6B showed low crossing-over frequency in the proximal regions. The distance from the centromere to Nor-B2 was only from 0.3 to 2.2 cM although it accounts for about two-thirds of the metaphase chromosome arm, which shows a great distortion of the metaphase map of the arm. The level of homologous recombination within the Nor-B2 locus is lower than in the chromosome region immediately distal to it. Whether it is comparable to that in the chromosome region proximal to it could not be determined. Recombination frequencies of different pairs of chromosome 6B in all but one interval paralleled the frequencies of their metaphase I pairing: Lower pairing at metaphase I was paralleled by lower crossing-over frequency. This relationship indicated that reduced metaphase I pairing between 6B chromosomes from different populations is due to impaired crossing-over and not due to precocious chiasma terminalization.     

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.     

Chromosomal location of genes for Rubisco small subunit and Rubisco-binding protein in common wheat

Summary The genes coding for the Rubisco small subunit (SSU) and for the -subunit of the Rubisco-binding protein were located to chromosome arms of common wheat. HindIII-digested total DNA from the hexaploid cultivar Chinese Spring and from ditelosomic and nullisomic-tetrasomic lines was probed with these two genes, whose chromosomal location was deduced from the disappearance of or from changes in the relative intensity of the relevant band(s). The Rubisco SSU pattern consisted of 14 bands, containing at least 21 different types of DNA fragments, which were allocated to two homoeologous groups: 15 to the short arm of group 2 chromosomes (4 to 2AS, 7 to 2BS, and 4 to 2DS) and 6 to the long arm of group 5 chromosomes (2 on each of arms 5AL, 5BL, and 5DL). The pattern of the Rubisco-binding protein consisted of three bands, each containing one type of fragment. These fragments were located to be on the short arm of group 2 chromosomes. The restriction fragment length polymorphism (RFLP) patterns of several hexaploid and tetraploid lines were highly conserved, whereas the patterns of several of their diploid progenitors were more variable. The variations found in the polyploid species were mainly confined to the B genome. The patterns of the diploids T. monococcum var. urartu and Ae. squarrosa were similar to those of the A and D genome, respectively, in polyploid wheats. The pattern of T. monococcum var. boeoticum was different from the patterns of the A genome, and the patterns of the diploids Ae. speltoides, Ae. longissima, and Ae. Searsii differed from that of the B genome.     

Influence of alien chromosomes on the resistance of soft wheat to biotrophic fungal pathogens

The resistance of 77 disomic chromosome-addition and chromosome-substitution lines of soft wheat that had been created on the base of the Chinese Spring variety to Belorussian populations of powdery mildew and brown leaf rust is studied. The lines possess a chromosome pair introduced from 13 species of cereals as well as three amphiploids. New genes that express resistance to biotrophic fungal pathogens in chromosomes 2S^s and 5S^s from the Aegilops searsii species and chromosome 6R^m from the Secale montanum species were discovered. We do not exclude the possibility that the new resistance gene against powdery mildew is carried by chromosome 6S¹ from the species Aegilops longissima (line No. 4), and that the new resistance gene against brown leaf rust is carried by chromosome 3E from the species Elytrigia elongata.     

普通小麦与鹅观草属间杂种F1及BC1的分子细胞遗传学、育性和赤霉病抗性研究

通过胚拯救, 成功获得鹅观草Roegneria kamoji (2n=6x=42, SSHHYY)和普通小麦中国春Triticum aesti-vum (2n=6x=42, AABBDD)的正反交属间杂种F1, 并对这些杂种F1及其BC1的形态学、减数分裂配对行为、育性和赤霉病抗性进行研究。结果表明, (鹅观草×中国春)F1和(中国春×鹅观草)F1的形态介于双亲之间。杂种F1花粉母细胞减数分裂中期I染色体构型分别为40.33I + 0.78II + 0.03III和40.40I + 0.79II 。杂种F1高度雄性不育, 用中国春花粉与其回交可获得BC1代种子。(鹅观草×中国春) F1×中国春BC1植株的染色体数目主要分布在55~63之间, 单价体较多, 植株高度不育; (中国春×鹅观草)F1×中国春BC₁植株染色体数目也主要分布在55~63之间, 但其中部分植株拥有整套小麦染色体且能正常配对、分离, 可形成部分可育花粉粒, 能收到少量自交结实种子。在 (鹅观草×中国春)F1中有1株穗型趋向中国春, 其染色体数目为2n=63, 经染色体分子原位杂交(GISH)检测, 含有42条小麦染色体和21条鹅观草染色体。该杂种F1在减数分裂中期I平均每个花粉母细胞有26.40I+18.30II, 但植株高度雄性不育, 用中国春花粉回交能收到BC₁种子。(鹅观草×中国春) F₁ (2n=63)×中国春BC₁的染色体数目主要分布在40~59之间, 其中的外源染色体已经逐渐减少, 虽然该BC₁的穗型已接近中国春, 但仍然高度不育。赤霉病抗性鉴定结果显示, 所有杂种F1及大部分BC₁对赤霉病均表现出较好的抗性。     

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.     

Substitutions of 2S and 7U chromosomes of Aegilops kotschyi in wheat enhance grain iron and zinc concentration

Biofortification through genetic manipulation is the best approach for improving micronutrient content of the staple food crops to alleviate hidden hunger, namely, the deficiency of Fe and Zn affecting more than two billion people worldwide. An interspecific hybridization was made between T. aestivum line Chinese Spring (CS) and Aegilops kotschyi accession 3790 selected for high grain iron and zinc concentration. The CS × Ae. kotschyi F₁ hybrid with low chromosome pairing was highly male and female sterile. This was backcrossed with wheat cultivars to get seed set. The selfed BC₁F₁ and BC₂F₁ plants with high grain iron and zinc concentration were selected in subsequent generations. The selected derivatives showed 60–136% enhanced grain iron and zinc concentration and 50–120% increased iron and zinc content per seed as compared to the recipient wheat cultivars. Thirteen cytologically stable, fertile and agronomically superior plants with high grain iron and zinc concentrations were selected for molecular characterization. The application of anchored wheat SSR markers, transferable to Ae. kotschyi, to the high grain iron and zinc containing derivatives indicated introgression of group 2 and group 7 chromosomes of Ae. kotschyi. GISH and FISH analysis of some derivatives confirmed the substitution of chromosomes 2S and 7U for their homoeologues of the A genome, suggesting that some of the genes controlling high grain micronutrient content in the Ae. kotschyi accession are on these chromosomes.     

Genetic compensation abilities of Aegilops speltoides chromosomes for homoeologous B-genome chromosomes of polyploid wheat in disomic S(B) chromosome substitution lines

The S genome of Aegilops speltoides is closely related to the B and G genomes of polyploid wheats. However, little work has been reported on the genetic relationships between the S-genome and B-genome chromosomes of polyploid wheat. Here, we report the isolation of a set of disomic substitutions (DS) of S-genome chromosomes for the B-genome chromosomes and their effects on gametophytic and sporophytic development. Ae. speltoides chromosomes were identified by their distinct C-banding and fluorescence in situ hybridization patterns with the Ae. speltoides-derived clone pGc1R-1. Although no large structural differences between S-genome and B-genome chromosomes exist, significant differences in gametophytic compensation were observed for chromosomes 1S, 3S, 5S and 6S. Similarly, chromosomes 1S, 2S, 4S, 5S and 6S affected certain aspects of sporophytic development in relation to spike morphology, fertility and meiotic pairing. The DS5S(5B) had disturbed meiosis with univalents/multivalents and suffered chromosome elimination in the germ tissues leading to haploid spikes in 50% of the plants. The effect of the Ph1 gene on meiosis is well known, and these results provide evidence for the role of Ph1 in the maintenance of polyploid genome integrity. These and other data are discussed in relation to the structural and functional differentiation of S- and B-genome chromosomes and the practical utility of the stocks in wheat improvement.