Genetic analysis of the species cytoplasm specific gene (scs d) derived from durum wheat

The action of species cytoplasm specific (scs) gene(s) can be observed when a durum (Triticum turgidum L.) nucleus is placed in the Aegilops longissimum S. & M. cytoplasm. This alloplasmic combination, (lo) durum, results in nonviable progeny. A scs gene derived from T. timopheevii Zhuk. (scs(ti)) produced compatibility with the (lo) cytoplasm. The resulting hemizygous (lo) scs(ti)- durum line was male sterile and when crossed to normal durum produced a 1:1 ratio of plump, viable (PV) seeds with scs(ti) and shriveled inviable (SIV) seeds without scs(ti). In a systematic characterization of durum lines an unusual line was identified that when crossed to (lo) scs(ti)- produced all PV seeds. When planted these PV seeds segregated at a 1:1 ratio of normal vigor plants (NVPs) and low vigor plants (LVPs). The LVP senescence before full maturity. The NVPs were male sterile and when crossed to common durum lines resulted in all plump seeds that again segregated at a 1:1 ratio of NVPs to LVPs. The crosses of these NVPs to common durum lines resulted in a 1:1 ratio of PV to SIV seeds. This study was extended to 317 individuals segregating for scs(ti) and the new locus, derived from durum wheat (scs(d)), establishing the allelic relationship of these two genes.     

Natural or induced nucleocytoplasmic heterogeneity in Triticum longissimum.

Alien cytoplasms produce a variety of phenotypes in durum wheat (Triticum turgidum) and common wheat (Triticum aestivum) cultivars, which indicate the prevalence of cytoplasmic variability in the subtribe Triticinae. Intraspecific cytoplasmic differences have been demonstrated between the subspecies of Triticum speltoides, Triticum dichasians, and Triticum comosum. In this study, durum wheat lines with cytoplasm from two accessions, B and C, of Triticum longissimum were compared, and meiotic chromosome pairing between the group 4 homoeologues from the same two accessions was examined in common wheat. First, monosomic addition or monosomic substitution lines of common wheat with cytoplasm and one chromosome (designated B) from accession B were crossed with those having cytoplasm and a chromosome designated C-1 or C-2 from accession C. In each substitution line, an alien chromosome substituted for a group 4 homoeologue. Each alien chromosome had a "selfish" (Sf) gene, which remained fixed in the wheat nucleus. The F1s had greatly reduced meiotic pairing between chromosomes B and C-1 and B and C-2, which indicated greatly reduced homology between the group 4 homoeologues from the two accessions. Second, by using Triticum timopheevii as a bridging species, chromosome B in a common wheat line was eliminated and an euploid durum line with cytoplasm from accession B was obtained. This line was fertile. In contrast, a similarly produced durum line with cytoplasm from accession C was male sterile and retained a species cytoplasm specific (scs) nuclear gene from T. timopheevii. In conclusion, nuclear and cytoplasmic heterogeneity pre-existed between accessions B and C and they represent varieties or incipient subspecies in T. longissimum. Alternatively, the Sf genes produced chromosomal heterogeneity and mutated cytoplasmic genes from one or both accessions. Key words : meiotic drive, selfish gene (Sf), gametocidal gene (Gc), Triticum, Aegilops.     

Isolation of a chromosomally engineered durum wheat line carrying the Aegilops ventricosa pchl gene for resistance to eyespot.

The chromosome 7Dv of Aegilops ventricosa (syn. Triticum ventricosum, 2n = 4x = 28, genome DvDvMvMv) carries the gene Pch1 for resistance to eyespot. This gene has previously been transferred to chromosome 7D of bread wheat, T. aestivum (2n = 6x = 42, genome AABBDD). To (1) enhance the level of resistance of bread wheat by increasing the copy number of Pch1, and (2) create eyespot-resistant triticales, meiotically stable Pch1-carrying durum lines were selected from the backcross progenies of a cross between Ae. ventricosa and T. durum cv. Creso ph1c (2n = 4x = 28, genome AABB). The Pch1 transfer, likely resulting from homoeologous recombination, was located at the distal position on the long arm of chromosome 7A. The 7A microsatellite marker Xgwm 698 was found closely linked in repulsion to the introgression in the resistant recombination lines, and the endopeptidase allele located on chromosome 7A of cv. Creso ph1c was lost.     

Detailed mapping of the species cytoplasm-specific (scs) gene in durum wheat

The compatibility-inducing action of the scs(ti) (species cytoplasm-specific gene derived from Triticum timopheevii) and Vi (vitality) genes can be observed when a durum (T. turgidum) nucleus is placed in T. longissimum cytoplasm. These two genes restore compatibility between an otherwise incompatible nucleus and cytoplasm. The objective of this study was to localize the scs(ti) gene on a linkage map of chromosome 1A, which could eventually be used to clone the gene. The mapping population consisted of 110 F2 individuals derived from crossing a Langdon-T. dicoccoides chromosome 1A substitution line with a euplasmic (normal cytoplasm) line homozygous for the scs(ti) gene. Through a series of testcrosses the genotypes of the 110 individuals were determined: 22 had two copies, 59 had one copy, and 29 had no copy of the scs(ti) gene. Data from RFLP, AFLP, and microsatellite analysis were used to create a linkage map. The flanking marker loci found for the scs(ti) gene were Xbcd12 and Xbcd1449-1A.2 with distances of 2.3 and 0.6 cM, respectively. Nearly 10% of individuals in this population were double recombinant for a genetic interval of <3 cM. A blistering phenotype reminiscent of the phenotype observed in maize brittle-1 mutable was also evident in these individuals. The higher frequency of double recombination within this region and seed-blistering phenotype could be an indication of a transposable element(s) in this locus.     

Synthesis and cytological characterization of trigeneric hybrids of durum wheat with and without Ph1.

Wild grasses in the tribe Triticeae, some in the primary or secondary gene pool of wheat, are excellent reservoirs of genes for superior agronomic traits, including resistance to various diseases. Thus, the diploid wheatgrasses Thinopyrum bessarabicum (Savul. and Rayss) A. Love (2n = 2x = 14; JJ genome) and Lophopyrum elongatum (Host) A. Love (2n = 2x = 14; EE genome) are important sources of genes for disease resistance, e.g., Fusarium head blight resistance that may be transferred to wheat. By crossing fertile amphidiploids (2n = 4x = 28; JJEE) developed from F1 hybrids of the 2 diploid species with appropriate genetic stocks of durum wheat, we synthesized trigeneric hybrids (2n = 4x = 28; ABJE) incorporating both the J and E genomes of the grass species with the durum genomes A and B. Trigeneric hybrids with and without the homoeologous-pairing suppressor gene, Ph1, were produced. In the absence of Ph1, the chances of genetic recombination between chromosomes of the 2 useful grass genomes (JE) and those of the durum genomes (AB) would be enhanced. Meiotic chromosome pairing was studied using both conventional staining and fluorescent genomic in situ hybridization (fl-GISH). As expected, the Ph1-intergeneric hybrids showed low chromosome pairing (23.86% of the complement), whereas the trigenerics with ph1b (49.49%) and those with their chromosome 5B replaced by 5D (49.09%) showed much higher pairing. The absence of Ph1 allowed pairing and, hence, genetic recombination between homoeologous chromosomes. Fl-GISH analysis afforded an excellent tool for studying the specificity of chromosome pairing: wheat with grass, wheat with wheat, or grass with grass. In the trigeneric hybrids that lacked chromosome 5B, and hence lacked the Ph1 gene, the wheat-grass pairing was elevated, i.e., 2.6 chiasmata per cell, a welcome feature from the breeding standpoint. Using Langdon 5D(5B) disomic substitution for making trigeneric hybrids should promote homoeologous pairing between durum and grass chromosomes and hence accelerate alien gene transfer into the durum genomes.     

Interactions between the scs and Vi genes in alloplasmic durum wheat.

Two nuclear genes, vitality (Vi) on an A- or B-genome chromosome and species cytoplasm specific (scs) on a 1DL telosome from Triticum aestivum L. or a telosome from Aegilops uniaristata Vis. (un telosome), improved compatibility between the nucleus of Triticum turgidum L. var. durum and the cytoplasm of Ae. longissima S. &M. or Ae. uniaristata. To study interactions between Vi and scs and to determine the chromosomal location of Vi, 29-chromosome fertile plants were crossed with 13 D-genome disomic-substitution (d-sub) lines [except 5D(5A)] of 'Langdon' durum. F1 and backcross progenies were examined for meiotic chromosome number and pairing, fertility, and plant vigor. In 11 crosses, Vi restored seed viability but produced double-monosomics (d-monos) with greatly reduced growth and vigor. In contrast, crosses involving 1D(1A) and 1D(1B) d-sub lines produced d-monos with normal vigor and anthesis but nonfunctional pollen. A backcross of 1D + 1A d-mono F1 and 1D(1A) d-sub lines produced 11 male steriles; 3 had 13 II + 1 II 1D + 1 I 1A, 2 had 13 II + 2 I, 1 had 13 II + 1 II 1D(1A), and 5 were not examined. Crosses of 1D + 1A d-mono F1 with control durum, lo durum (with 1DL), and un durum (with un telosome) lines produced 16 male-sterile d-monos and 14 fertiles with 14 II + 1 I 1D, showing that 15-chromosome female gametes transmitted monosomes 1A and 1D. However, BC2F1's from 1D + 1B d-mono x fertile line with un telosome included 20 male-sterile d-monos, 6 fertile triple monosomics (13 II + 1 I 1D + 1 I 1B + t I un telosome), and 1 fertile plant with a 1B/1D translocation. Unlike d-mono 1A + 1D, d-mono 1B + 1D did not transmit 15-chromosome female gametes with monosomes 1D and 1B. Additional backcrosses also indicated that homozygous scs caused male sterility in 1D(1A) and 1D(1B) d-subs and that the procedure used was not suitable for the chromosomal location of Vi.     

Cytological and molecular characterization of a Triticum aestivum x Lophopyrum ponticum backcross derivative resistant to barley yellow dwarf.

Barley yellow dwarf is the most damaging virus-caused disease in bread wheat (Triticum aestivum L.). A resistant line, SW335.1.2-13-11-1-5 (2n = 47), derived from a cross of T. aestivum x Lophopyrum ponticum was characterized by meiotic chromosome pairing, by in situ DNA hybridization and by expression of molecular markers to determine its chromosome constitution. All progeny of this line had three pairs of L. ponticum chromosomes from homoeologous chromosome groups 3, 5, and 6 and the 2n = 47 progeny had an additional L. ponticum monosome. The pairs from groups 3 and 6 were in the added state, while the group 5 pair was substituted for wheat chromosome 5D. Several wheat-wheat translocations with respect to the parental wheat genotype occurred in this line, presumably owing to the promotion of homoeologous chromosome pairing by L. ponticum chromosomes. It was hypothesized that homoeologous recombination results in homoeologous duplication-deletions in wheat chromosomes. An aberrant 3:1 disjunction creates the potential at each meiosis for replacement of these wheat chromosomes by homoeologous L. ponticum chromosomes. Wheat chromosomes 3A and 6A appeared to be in intermediate stages of this substitution process.     

A fertile amphiploid between durum wheat (Triticum turgidum) and the x Agroticum amphiploid (Agropyron cristatum x T. tauschii)

Agropyron (Gaertn) is a genus of Triticeae which includes the crested wheatgrass complex, i.e. A. cristatum (L.) as representative species containing the P genome. This species is an important source for increase the genetic variability of both durum and bread wheat. Among the possible interesting features to be introgressed into wheat are resistance to wheat streak mosaic virus, rust diseases, and tolerance to drought, cold and moderate salinity. By crossing tetraploid wheat (Triticum turgidum conv durum, 2n = 4x = 28; AABB) with a fertile allotetraploid (2n = 4x = 28; DDPP) between diploid wheat (T. tauschii) and crested wheatgrass (A. cristatum L.), amphiploid plants were obtained. Fluorescence in situ hybridization (FISH) using both genomic DNA from A. cristatum and the repetitive probe pAs1, proved that the plants were true amphiploids with a chromosome number 2n = 8x = 56 and genomic constitution AABBDDPP. Using total genomic in situ hybridization (GISH) to study meiotic metaphase I, data on allosyndetic and autosyndetic chromosome pairing were obtained. The amphiploids were perennial like the male parent but their morphology was close to that of the wheat parent. They were resistant to wheat leaf rust and powdery mildew under field conditions.     

Standard karyotype of Triticum longissimum and its cytogenetic relationship with T. aestivum.

C-banding polymorphism was analyzed in 17 accessions of Triticum longissimum from Israel and Jordan, and a generalized idiogram of this species was established. C-banding analysis was further used to identify two sets of disomic T. aestivum - T. longissimum chromosome addition lines and 13 ditelosomic addition lines and one monotelosomic (6S1L) addition line. C-banding was also used to identify T. aestivum - T. longissimum chromosome substitution and translocation lines. Two major nucleolus organizing regions (NORs) on 5S1 and 6S1 and one minor NOR on 1S1 were detected by in situ hybridization using a 18S-26S rDNA probe. Sporophytic and gametophytic compensation tests were used to determine the homoeologous relationships of T. longissimum chromosomes. The T. longissimum chromosomes compensate rather well and fertility was restored even in substitution lines involving wheat chromosomes 2A, 4B, and 6B that contain major fertility genes. Except for the deleterious gametocidal genes, T. longissimum can be considered as a suitable donor of useful genes for wheat improvement.     

Interspecific nuclear-cytoplasmic compatibility controlled by genes on group 1 chromosomes in durum wheat.

Triticum longissimum cytoplasm is incompatible with the T. turgidum nuclear genome. Two nuclear genes, scs and Vi, derived from the nuclear genome of T. timopheevii and by a spontaneous mutation, respectively, restore nuclear-cytoplasmic compatibility, normal plant vigor, and male fertility in these alloplasmic genotypes. The objectives of this study were (i) to determine the chromosomal locations of scs and Vi; (ii) to identify DNA markers for scs and Vi; and (iii) to determine the interactions involving the dosage of scs and Vi. Two populations segregating for scs and Vi were produced and scored for seedling vigor (indicating presence of scs) and degree of self-fertility (indicating presence of Vi). Four RFLP markers were mapped near scs. Aneuploid analysis revealed that these markers, and hence the scs gene, are located on the long arm of chromosome 1A. Four RFLP markers were mapped near Vi on 1BS. Results indicated that other factors may be inhibiting the expression of Vi. We determined the dosage of scs and Vi in both populations with the aid of the linked RFLP markers. Individuals with two versus one dose of scs had reduced self-fertility, while individuals with two versus one dose of Vi had similar self-fertility.     

Radiation hybrid mapping of the species cytoplasm-specific (scsae) gene in wheat

Radiation hybrid (RH) mapping is based on radiation-induced chromosome breakage and analysis of chromosome segment retention or loss using molecular markers. In durum wheat (Triticum turgidum L., AABB), an alloplasmic durum line [(lo) durum] has been identified with chromosome 1D of T. aestivum L. (AABBDD) carrying the species cytoplasm-specific (scsae) gene. The chromosome 1D of this line segregates as a whole without recombination, precluding the use of conventional genome mapping. A radiation hybrid mapping population was developed from a hemizygous (lo) scsae--line using 35 krad gamma rays. The analysis of 87 individuals of this population with 39 molecular markers mapped on chromosome 1D revealed 88 radiation-induced breaks in this chromosome. This number of chromosome 1D breaks is eight times higher than the number of previously identified breaks and should result in a 10-fold increase in mapping resolution compared to what was previously possible. The analysis of molecular marker retention in our radiation hybrid mapping panel allowed the localization of scsae and 8 linked markers on the long arm of chromosome 1D. This constitutes the first report of using RH mapping to localize a gene in wheat and illustrates that this approach is feasible in a species with a large complex genome.     

Comparison of homoeologous group-6 short arm physical maps of wheat and barley reveals a similar distribution of recombinogenic and gene-rich regions

Eighty two new loci, mapped with 51 DNA clones, were added to the earlier deletion maps of the homoeologous group-6 short arms of hexaploid wheat ( Triticum aestivum L. em Thell., 2n = 6 x = 42, AABBDD). There are now 41, 56 and 52 loci mapped on deletion maps of 6AS, 6BS and 6DS, respectively. The linear order of orthologous loci in all three arms appears to be identical. The majority of the loci are located in the distal one-half of the three arms. There seems to be an increased marker/gene density from the centromeric to the telomeric regions in each arm, and the marker density in comparable physical regions is similar on all three maps. Recombination is not uniformly distributed along the chromosome arms; 60% of recombination occurs in the distal one-third of each arm. Recombination increases from the proximal region to the distal end in a nonlinear pattern. The distribution of loci and recombination along each of the three chromosome arms is highly correlated. Comparison of the 6BS deletion map from this study and a 6HS physical map of barley ( Hordeum vulgare L., 2n = 2 x = 14, HH) reveals a remarkably similar distribution of recombinogenic and gene-rich regions between the two chromosome arms, suggesting that the distribution patterns of genes may be conserved in the homoeologous group-6 chromosome short arms of wheat and barley. A consensus map of wheat group-6 short arms containing 46 orthologous loci was constructed. Comparison of the consensus map with published linkage maps of Triticeae group-6 chromosome short arms indicates that the linear order of the loci on the maps has been largely conserved. Evidence from this study does not support the existence of a 2BS-6BS reciprocal terminal translocation.     

C-banding and fluorescence in situ hybridization studies of the wheat-alien hybrid 'Agrotana'.

'Agrotana', a wheat-alien hybrid (2n = 56), is a potential source of resistance to common root rot, stem rust, wheat streak mosaic virus, and the wheat curl mite. However, the origin of 'Agrotana', reported to be durum wheat x Agropyron trichophorum (pubescent wheatgrass), is uncertain. The objective of this investigation was to determine the chromosome constitution of 'Agrotana' using C-banding and fluorescence in situ hybridization techniques. The F1 hybrid of 'Agrotana' x 'Chinese Spring' wheat showed 7 I + 21 II in 14.9% of the pollen mother cells, evidence of the presence of the A, B, and D genomes in 'Agrotana'. The hybrid had 16 heavily C-banded chromosomes, namely 4A, and 1-7B of wheat, and a translocation that probably involved wheat chromosomes 2A and 2D. In situ hybridization using biotinylated genomic DNA of Ag. trichophorum cv. Greenleaf blocked with CS DNA failed to identify the alien chromosomes in 'Agrotana', indicating that the alien chromosomes were not likely derived from pubescent wheatgrass. In situ hybridization using labelled wheat genomic DNA blocked with 'Agrotana' DNA revealed that 'Agrotana' had 40 wheat, 14 alien, and 2 (a pair) wheat-alien translocated chromosomes. There was no homology between wheat and the alien chromosomes or chromosome segments involved in the wheat-alien recombinant. Two of the seven pairs of alien chromosomes were homoeologous to each other. The ability to identify alien chromatin in wheat using labelled wheat DNA instead of labelled alien DNA will be particularly useful in chromosome engineering of wheat germplasms having alien chromatin of unknown origin.     

Molecular cytogenetic analysis of durum wheat x tritordeum hybrids.

Southern and in situ hybridization were used to examine the chromosome constitution, genomic relationships, repetitive DNA sequences, and nuclear architecture in durum wheat x tritordeum hybrids (2n = 5x = 35), where tritordeum is the fertile amphiploid (2n = 6x = 42) between Hordeum chilense and durum wheat. Using in situ hybridization, H. chilense total genomic DNA hybridized strongly to the H. chilense chromosomes and weakly to the wheat chromosomes, which showed some strongly labelled bands. pHcKB6, a cloned repetitive sequence isolated from H. chilense, enabled the unequivocal identification of each H. chilense chromosome at metaphase. Analysis of chromosome disposition in prophase nuclei, using the same probes, showed that the chromosomes of H. chilense origin were in individual domains with only limited intermixing with chromosomes of wheat origin. Six major sites of 18S-26S rDNA genes were detected on the chromosomes of the hybrids. Hybridization to Southern transfers of restriction enzyme digests using genomic DNA showed some variants of tandem repeats, perhaps owing to methylation. Both techniques gave complementary information, extending that available from phenotypic, chromosome morphology, or isozyme analysis, and perhaps are useful for following chromosomes or chromosome segments during further crossing of the lines in plant breeding programs.     

不同细胞质的普通小麦"中国春"(Triticum aestivum var.Chinese Spring)与小偃麦杂交F1代的育性与减数分裂行为的研究

15个不同细胞质“中国春”小麦与八倍体小偃麦杂交 ,杂种F1减数分裂的染色体行为表明 :普通小麦与天蓝偃麦草的F或E组染色体之间存在着部分同源关系 ;D2 型细胞质促进部分同源染色体配对、但却抑制同源染色体配对 ;Sv 型细胞质对同源染色体或部分同源染色体的配对均有抑制作用 ;G型细胞质促进同源染色体配对。1 5个不同细胞质“中国春”小麦与六倍体小偃麦杂交 ,F1结实率很低 ,减数分裂中期的染色体行为混乱 ,单价体过多 ,或许意味着在天蓝偃麦草 (Elytrigiain termedium)与长穗偃麦草 (E .elongatum)的E组染色体之间存在着很大差别。随着回交代数的增加 ,选出G型、D2 型、Mt 型、Mu 型等细胞质雄性不育的八倍体小偃麦品系 ,其中D2 型细胞质八倍体小偃麦具有光周期敏感性雄性不育的特征 ;G型细胞质“远中 3”育性正常 ,表明八倍体小偃麦“远中 3”的E组染色体中存在G型胞质的育性恢复基因。     

Chromosome sorting in tetraploid wheat and its potential for genome analysis

This study evaluates the potential of flow cytometry for chromosome sorting in durum wheat (Triticum turgidum Desf. var. durum, 2n = 4x = 28). Histograms of fluorescence intensity (flow karyotypes) obtained after the analysis of DAPI-stained chromosomes consisted of three peaks. Of these, one represented chromosome 3B, a small peak corresponded to chromosomes 1A and 6A, and a large peak represented the remaining 11 chromosomes. Chromosomes sorted onto microscope slides were identified after fluorescence in situ hybridization (FISH) with probes for GAA microsatellite, pSc119.2, and Afa repeats. Genomic distribution of these sequences was determined for the first time in durum wheat and a molecular karyotype has been developed for this crop. Flow karyotyping in double-ditelosomic lines of durum wheat revealed that the lines facilitated sorting of any arm of the wheat A- and B-genome chromosomes. Compared to hexaploid wheat, flow karyotype of durum wheat is less complex. This property results in better discrimination of telosomes and high purities in sorted fractions, ranging from 90 to 98%. We have demonstrated that large insert libraries can be created from DNA purified using flow cytometry. This study considerably expands the potential of flow cytogenetics for use in wheat genomics and opens the possibility of sequencing the genome of this important crop one chromosome arm at a time.     

Meiotic pairing in hybrids between tetraploid Triticale and related species: new elements concerning the chromosome constitution of tetraploid Triticale

Summary Two F5 strains of tetraploid triticale (2n= 4x=28), obtained from 6x triticaleX2 rye progenies, were crossed with diploid and tetraploid rye, some durum and bread wheats, and various 8x and 6x triticale lines. Meiosis in the different hybrid combinations was studied. The results showed that the haploid complement of these triticales consists of seven chromosomes from rye and seven chromosomes from wheat. High frequencies of PMCs showing trivalents were observed in hybrids involving the reference genotypes of wheat and triticale. These findings proved that several chromosomes from the wheat component have chromosome segments coming from two parental wheat chromosomes. The origin of these heterogeneous chromosomes probably lies in homoeologous pairing occurring at meiosis in the 6x triticaleX2x rye hybrids from which 4x triticale lines were isolated. A comparison among different hybrids combinations indicated that the involvement of D-genome chromosomes in homoeologous pairing is quite limited. In contrast, meiotic patterns in 4x triticale X 2x rye hybrids showed a quite high pairing frequency between some R chromosomes and their A and B homoeologues.     

Chromosomal locations of the genes for histones and a histone gene-binding protein family HBP-1 in common wheat

The chromosomal locations of the genes in common wheat that encode the five histones and five members of the HBP (histone gene-binding protein)-1 family were determined by hybridizing their cloned DNAs to genomic DNAs of nullitetrasomic and telosomic lines of common wheat, Triticum aestivum cv. Chinese Spring. The H1 and H2a genes are located on different sets of homoeologous chromosomes or chromosome arms, namely, 5A, 5B and 5D, and 2AS, 2BS and 2DS, respectively. Genes for the other histones, H2b, H3 and H4, are found in high copy number and are dispersed among a large number of chromosomes. The genes for all members of the HBP-1 family are present in small copy numbers. Those for HBP-1a(1) are located on six chromosome arms, 3BL, 5AL, 5DL, 6AL, 6BS and 7DL, whereas those for each HBP-1a(c14), 1a(17), 1b(c1), and 1b(c38) are on a single set of homoeologous chromosome arms; 4AS, 4BL, 4DL; 6AS, 6BS, 6DS; 3AL, 3BL, 3DL; and 3AS, 3BS, 3DS, respectively. The genes for histones H1 and H2a, and for all members of the HBP-1 family except HBP-1a(1) are assumed to have different phylogenetic origins. The genes for histone 2a and HBP-1a(17) are located in the RFLP maps of chromosomes 2B and 6A, respectively. Gene symbols are proposed for all genes whose chromosomal locations have been determined.     

Chromosome engineering in wheat to restore male fertility in the msH1 CMS system

Pollen fertility restoration of the CMS phenotype caused by H. chilense cytoplasm in wheat was associated with the addition of chromosome 6H^chS from H. chilense accession H1. In order to develop an euploid restored line, different genomic combinations substituting the 6H^chS arm for another homoeologous chromosome in wheat were evaluated, with the conclusion that the optimal combination was the translocation T6H^chS·6DL. The double translocation T6H^chS·6DL in H. chilense cytoplasm was obtained. This line is fertile and stable under different environmental conditions. However, a single dose of the T6H^chS·6DL translocation is insufficient for fertility restoration when chromosome 6D is also present. Restoration in the msH1 system is promoted by interaction between two or more genes, and in addition to the restorer of fertility (Rf) located on chromosome 6H^chS, one or more inhibitor of fertility (Fi) genes may be present in chromosome 6DL.     

Microsatellite mapping of the genes for brittle rachis on homoeologous group 3 chromosomes in tetraploid and hexaploid wheats

The brittle rachis character, which causes spontaneous shattering of spikelets, has an adaptive value in wild grass species. The loci Br1 and Br2 in durum wheat (Triticum durum Desf.) and Br3 in hexaploid wheat (T. aestivum L.) determine disarticulation of rachides above the junction of the rachilla with the rachis such that a fragment of rachis is attached below each spikelet. Using microsatellite markers, the loci Br1, Br2 and Br3 were mapped on the homoeologous group 3 chromosomes. The Br2 locus was located on the short arm of chromosome 3A and linked with the centromeric marker, Xgwm32, at a distance of 13.3 cM. The Br3 locus was located on the short arm of chromosome 3B and linked with the centromeric marker, Xgwm72 (at a distance of 14.2 cM). The Br1 locus was located on the short arm of chromosome 3D. The distance of Br1 from the centromeric marker Xgdm72 was 25.3 cM. Mapping the Br1, Br2 and Br3 loci of the brittle rachis suggests the homoeologous origin of these 3 loci for brittle rachides. Since the genes for brittle rachis have been retained in the gene pool of durum wheat, the more closely linked markers with the brittle rachis locus are required to select against brittle rachis genotypes and then to avoid yield loss in improved cultivars.