Rye Chromosome 1R Research Articles

The production of a 5B-5R chromosome translocation for induction of homoeologous meiotic pairing in wheat

A wheat line heterozygous for a translocation between the short arm of wheat chromosome 5B and the long arm of rye chromosome 5R was isolated. The rye segment carries the gene Hp, which gives the plants hairy peduncles. The rye segment does not carry the homoeologous pairing suppressor gene Ph 1 of the long arm of the 5B chromosome of wheat. This makes the translocation line very useful for the induction of homoeologous chromosome pairing between alien chromosomes and wheat chromosomes in interspecific and intergeneric hybrids. Such hybrids with this translocation line will be of two kinds: with or without the Hp gene. The former lack the Phi gene and should have homoeologous chromosome pairing in meiosis. The production of the translocation line is described.

Dissection of rye chromosome 1R in common wheat

Rye chromosome 1R contains many agronomically useful genes. Physical dissection of chromosome 1R into segments would be useful in mapping 1R-specific DNA markers and in assembling DNA clones into contig maps. We applied the gametocidal system to produce rearranged 1R chromosomes of Imperial rye (1R(i)) added to common wheat. We identified rearranged 1R(i) chromosomes and established 55 1R(i) dissection lines of common wheat carrying a single rearranged 1R(i) chromosome. Fifty-two of the rearranged 1R(i) chromosomes had single breakpoints and three had double breakpoints. The 58 breakpoints were distributed in the short arm excluding the satellite (12 breakpoints), in the satellite (4), in the long arm (28), and in the centromere (14). Out of the 55 lines, nine were homozygous for the rearranged 1R(i) chromosomes, and the remaining lines were hemizygous. We developed 26 PCR-based EST markers that were specific to the 1R(i) chromosome, and nine of them amplified 1R(i) arm-specific PCR products without restriction-enzyme digestion. Using the nine EST markers and two previously reported 1R-specific markers, we characterized the 55 1R(i) dissection lines, and also proved that we can select critical progeny plants carrying specific rearranged 1R(i) chromosomes by PCR, without cytological screening, in 48 out of the 55 hemizygous dissection lines.

Genetic regulation of the centromere division in rye and wheat univalent chromosomes in dimonosomics during the meiotic anaphase I

Cytogenetic analysis of meiosis in the wheat--rye dimonosomics 1Rv-1A, 1Ron-1A, 2R-2D, 5R-5A, and 6R-6A was conducted. C-banding was used to study the segregation pattern of each of two univalent chromosomes during the first meiotic division. It has been shown that the division frequency of the centromeric regions of all rye chromosomes in the pair studied is significantly higher than in the wheat chromosomes. The ANOVA performed suggest that the plant genotype contributes significantly (at P = 0.05) to the behavior pattern of univalent chromosomes in meiosis. The data obtained demonstrate that the rye and wheat chromosomes studied are involved in genetic regulation of centromere division in meiotic anaphase I (AI). The presence of rye chromosome 2R and wheat chromosome 2D suppresses the division of centromeres of the sister chromatids in AI. Rye chromosomes 1Rv, 1Ron, 5R, and 6R induce equational division; however, rye chromosome 1Rv increases to a greater degree the frequency of equational division of wheat chromosome 1A as compared with chromosome 1Ron.

Gene Expression and Nucleolar Dominance in Hexaploid Triticale and T. Aestivum

ABSTRACTThe expression of rRNA genes located in the nucleolar organizing region (NOR) present in the short arm of chromosome 1R from rye (Secale cereale L.) was examined in several hexaploid genotypes of T. aestivum cv. Chinese Spring (AABBDD), Triticale 171 (AABBRR), sphaerococcum mutant form MT47–2D(2R) and Secale cereale (RR). By silver staining and in situ (FISH) hybridization with the DNA probe pTa71, was established a corresponance between the expression and organization patterns of rDNA domains in metaphase and interphase cells. We show here that the substitution of the rye chromosome 2R by a chromosome from hexaploide wheat 2D (triticale–2D(2R) in MT47sph, prevents suppression of the rye-origin rDNA and leads to activity of all six major rDNA loci. In spite of that in triticale 171, containing 28 chromosomes of wheat origin and 14 from rye, rDNA of rye origin is not normally expressed, while the 1B- and 6B- origin rDNA from wheat shows strong expression.

Estimating meiotic chromosome pairing and recombination parameters in telocentric trisomics

Telocentric trisomics (telotrisomics; one arm of a metacentric chromosome present in addition to two complete genomes) are used in theoretical studies of pairing affinities and chiasma formation in competitive situations and applied in genome analysis, gene localization, gene transfer, and breakage of close linkages. These applications require knowledge of the recombination characteristics of telotrisomics. Appropriate cytological and molecular markers and favorable chromosome morphology are not always available or applicable for quantitative analyses. We developed new mathematical models for extracting the maximum information from simple metaphase I observations. Two types of telotrisomics of the short arm of chromosome 1R of rye (Secale cereale), including several genotypes, were used as test material. In simple telotrisomics, pairing between morphologically identical complete chromosomes was more frequent than pairing between the telocentric and either of the normal chromosomes. In the telocentric substitution, morphologically identical telocentrics paired less frequently with each other than either one with the normal chromosome. Pairing partner switch was significant. Interaction between the two arms was variable. Variation within plants was considerable. Telotrisomics without markers are suitable for analyzing pairing preferences, for gene localization and gene transfer, and for breaking tight linkages, but less so for genome analysis.

Molecular Mapping of the Russian Wheat Aphid Resistance Gene Dn2414 in Wheat

Russian wheat aphid (RWA) [Diuraphis noxia (Kurdjumov)] is an important pest of wheat (Triticum aestivum L.) in several production areas of the world. The most effective and economical approach for controlling RWA is to use resistant cultivars. A wheat line, ST‐ARS 02RWA2414‐11 (2414‐11), showed a high level of resistance to RWA biotype 2. Our objectives were to map the resistance gene and develop polymerase chain reaction (PCR)–based markers for marker‐assisted selection (MAS). A mapping population of 212 F2 individuals was developed from a cross of 2414‐11 and the susceptible cultivar Yuma. The F2 individuals and F2:3 families were infested using biotype 2 RWA. The RWA resistance of 2414‐11 is controlled by a single major gene, provisionally designated as Dn2414 Using standard PCR, 30 marker loci were found to be linked to Dn2414 with recombination frequencies (θ) of 0.00 to 0.27 and logarithm of the odds to the base 10 (LOD) scores of 7.6 to 66.1. Of the 30 markers, 26 were tightly linked to Dn2414 with θ ≤ 0.05. A genetic map was constructed consisting of 31 loci spanning a region of 34.7 cM. The close linkage of Dn2414 with several rye chromosome 1R short arm (1RS)‐specific simple sequence repeat markers and low θ values around the Dn2414 gene indicate that Dn2414 is located on chromosome 1RS.1BL (translocation chromosome with IRS and wheat chromosome 1B long arm). Phenotypic and marker profiles of 2414‐11 and its relatives are the same as other lines known to carry Dn7 The Dn2414 gene is thus located on 1RS arm, and the large number of PCR markers will be valuable for MAS of this gene.

Effect of rye Secale cereale L. chromosomes 1R and 3R on polyembryony expression in hybrid combinations between (Hordeum L.)-Triticum aestivum L. alloplasmic recombinant lines and wheat-rye substitution lines T. aestivum L.-S. cereale L.

The effect of rye chromosomes on polyembryony was studied for reciprocal hybrid combinations between (Hordeum vulgare L.)-Triticum aestivum L. alloplasmic recombinant lines and five wheat T. aestivum L. (cultivar Saratovskaya 29)-rye Secale cereale L. (cultivar Onokhosikaya) substitution lines: 1R(1D), 2R(2D), 3R(3B), 5R(5A), and 6R(6A), and for direct hybrid combinations between the [H. marinum ssp. gussoneanum (H. geniculatum All.)]-T. aestivum alloplasmic recombinant line and the wheat-rye substitution lines 1R(1A), 1R(1D), and 3R(3B). Chromosomes 1R and 3R of rye cultivar Onokhoiskaya proved to affect the expression of polyembryony in the hybrid combinations that involved the alloplasmic recombinant lines of common wheat as maternal genotypes. Based on this finding, polyembryony was regarded as a phenotypic expression of nuclear-cytoplasmic interactions where an important role is played by rye chromosomes 1R and 3R and the H. vulgare cytoplasm. Consideration is given to the association between the effect of rye chromosomes 1R and 3R on polyembryony in the [(Hordeum)-T. aestivum × wheat-rye substitution lines] hybrid combinations and their stimulating effect on the development on androgenic embryoids in isolated anther cultures of the wheat-rye substitution lines.

Role of rye chromosome 2R from wheat-rye substitution line 2R(2D)1 (Triticum aestivum L. cv. Saratovskaya 29-Secale cereale L. cv. Onokhoiskaya) in genetic regulation of meiotic restitution in wheat-rye polyhaploids

A study was made of the role of rye chromosome 2R from the wheat-rye substitution line 2R(2D)1 (Triticum aestivum L. cv. Saratovskaya 29-Secale cereale L. cv. Onokhoiskaya) in genetic regulation of meiotic restitution in wheat-rye polyhaploids 2R(2D)1 x S. cereale L. cv. Onokhoiskaya. Rye chromosome 2R proved to affect the completeness of the meiotic program, suppressing the formation of restitution gametes. This was evident from the reductional division of univalent chromosomes in AI and the occurrence of the second meiotic division. The interrelationships between the type of chromosome division in AI and the two-step character of meiosis are discussed. The structural and functional organization of the centromeric regions of chromosomes undergoing reductional division is assumed to determine the two-step character of division.

Complex genome rearrangements reveal evolutionary dynamics of pericentromeric regions in the Triticeae

Most pericentromeric regions of eukaryotic chromosomes are heterochromatic and are the most rapidly evolving regions of complex genomes. The closely related genomes within hexaploid wheat (Triticum aestivum L., 2n=6x=42, AABBDD), as well as in the related Triticeae taxa, share large conserved chromosome segments and provide a good model for the study of the evolution of pericentromeric regions. Here we report on the comparative analysis of pericentric inversions in the Triticeae, including Triticum aestivum, Aegilops speltoides, Ae. longissima, Ae. searsii, Hordeum vulgare, Secale cereale, and Agropyron elongatum. Previously, 4 pericentric inversions were identified in the hexaploid wheat cultivar 'Chinese Spring' ('CS') involving chromosomes 2B, 4A, 4B, and 5A. In the present study, 2 additional pericentric inversions were detected in chromosomes 3B and 6B of 'CS' wheat. Only the 3B inversion pre-existed in chromosome 3S, 3Sl, and 3Ss of Aegilops species of the Sitopsis section, the remaining inversions occurring after wheat polyploidization. The translocation T2BS/6BS previously reported in 'CS' was detected in the hexaploid variety 'Wichita' but not in other species of the Triticeae. It appears that the B genome is more prone to genome rearrangements than are the A and D genomes. Five different pericentric inversions were detected in rye chromosomes 3R and 4R, 4Sl of Ae. longissima, 4H of barley, and 6E of Ag. elongatum. This indicates that pericentric regions in the Triticeae, especially those of group 4 chromosomes, are undergoing rapid and recurrent rearrangements.

Cytogenetically Engineered Rye Chromosomes 1R to Improve Bread‐making Quality of Hexaploid Triticale

Hexaploid triticale (X Triticosecale Wittmack) is rarely used for human consumption because of its poor bread‐making quality. To create the genetic potential for bread‐making quality similar to that of bread wheat (Triticum aestivum L), rye (Secale cereale L.) chromosome 1R in triticale cv. Presto was cytogenetically engineered to remove secalin loci Sec‐1 and Sec‐3, and to introduce wheat storage protein loci Gli‐1 and Glu‐1 The manipulations were by homoeologous recombination between rye chromosome 1R and 1B or 1D of wheat, followed by homologous recombination of primary recombinants with translocation breakpoints in desired locations. This approach generated three classes of multi‐breakpoint translocation chromosomes named Valdy, FC and RM. Chromosome Valdy is a three breakpoint translocation with loci Gli‐D1, Sec‐1 and Glu‐D1; chromosomes FC1 and FC2 are five breakpoint translocations with loci Gli‐D1 and Glu‐D1, and chromosomes RM are six breakpoint translocations with loci Gli‐B1 and Glu‐D1 Preliminary tests of the effects of these chromosomes show a 230 to 250% increase of the SDS‐sedimentation value over the recipient triticale Presto. While the impact of these chromosomes on the agronomic value of triticale is not clear, they restore to triticale the genetic load of gluten‐encoding loci similar to that of bread wheat, thereby creating the genetic potential for breeding bread‐making triticales.

Visualization of Secale Cereale DNA in Wheat Germ Plasm by Genomic In Situ Hybridization

ABSTRACTTotal genomic DNA from rye was labelled with biotin and used as a probe for in situ hybridization to show the sizes and translocation points of the rye chromosome segments in wheat mutant forms K1 and K2, which carry a translocation between wheat chromosome 1B, and the short arm of rye chromosome 1R (1B/1R). The translocation breakpoints were at or very near to, the centromere. Total genomic rye DNA used as a probe was also able to distinguish 12 rye chromosomes in 6x triticale-MT47 by genomic in situ hybridization. The techniques using genomic probes are useful for detecting, characterizing and following alien chromosomes or chromosome segments by genomic hybridization- GISH.

Transfer of a Rye Small Chromosomal Segment with Powdery Mil-dew- Resistant Gene(s) into Common Wheat (Triticum aestivum L.)

The advanced progeny lines (BC1F5) from the monosomic addition lines between common wheat cultivar Mianyang 11, which is highly susceptible to powdery mildew, and an inbred rye line R12 were analyzed for selection of wheat-rye translocations. Based on a rye-specific repetitive sequence of pSc20H, which spread over all chromosomes of rye but did not existed in wheat, a set of PCR primer was designed and used to identify the rye chromosome segments in wheat. From 300 of the BC1F5 progeny lines 70 were found to contain chromosome composition of rye. An advanced line, 96II691-830-98, originated from 6R monosomic addition line was observed to be immune to powdery mildew, different from its wheat parent Mianyang 11. A small segment of rye chromosome at telomere in a pair of wheat chromosome in the line was found by means of GISH. The results indicated that a small segment of rye chromosome 6R carrying the gene(s) for resistance to powdery mildew has been transferred into common wheat. In the progeny of monosomic addition lines a high frequency of wheat-alien species translocation with various segments of chromosomes could be found by application of both PCR and GISH technique.

Features of Crossability, Haploidy and Polyembryony in Hybrid Combinations between Cultivated Barley Hordeum vulgare L. (2n = 14) and Wheat-Rye Substitution Lines Triticum aestivum L., Cultivar Saratovskaya 29/Secale cereale L., Cultivar Onokhoiskaya

The role of individual chromosomes of rye in the manifestation of crossability and seedling development in hybrid combinations between common barley Hordeum vulgare L., cultivar Nepolegayushchii (2n = 14) and five wheat-rye substitution lines Triticum aestivum L., cultivar Saratovskaya 29/Secale cereale L., cultivar Onokhoiskaya (2n = 40 wheat + 2 rye chromosomes). Crossability, which was measured by two parameters--frequency of set grains and frequency of grains with embryos--was shown to be significantly affected by each of the five rye chromosomes examined: 1R, 2R, 3R, 5R, and 6R; the development of barley haploids was affected by rye chromosomes 1 R, 3R, and 5R. We were the first to demonstrate that polyembryony could be induced by mutual effects of barley cytoplasm and rye chromosome 1R. Possible mechanisms controlling the development of haploids and twins in hybrid combinations H. vulgare x T. aestivum/S. cereale are discussed. The conclusion is drawn that hybrid combinations between common barley and wheat-rye substitution lines can serve as new models for studying incompatibility mechanisms in distant crosses and genetic control of parthenogenesis.

Effect of the 5R(5A) Alien Chromosome Substitution on the Growth Habit and Winter Hardiness of Wheat

The growth habit, ear emergence time, and frost tolerance of wheat/rye substitution lines have been studied in cultivars Rang and Mironovskaya Krupnozernaya whose chromosome 5A is substituted with chromosome 5R of Onkhoyskaya rye. Hybrid analysis has demonstrated that the spring habit of the recipient cultivars Rang and Mironovskaya Krupnozernaya is controlled by dominant gene Vrn-A1 located in chromosome 5A. Onokhoyskaya rye has a dominant gene for the spring habit (Sp1) located in chromosome 5R. It has been found that the resultant 5R(5A) alien-substitution lines have a winter type of development and ears do not emerge during summer in plants sown in spring. The change in growth habit has been shown to be related to the absence of the rye Spl gene expression in the substitution lines. The winter hardiness of winter 5R(5A) alien-substitution lines has been studied under the environmental conditions of Novosibirsk. Testing the lines in the first winter demonstrated that their winter survival is 20-27%. The possible presence of the frost resistance gene homeoallelic to the known genes Fr1 and Fr2 of the common wheat located on chromosomes 5A and 5D, respectively, is discussed.

Chromosomal Assignment of the Genes Encoding Glutamyl-tRNA Reductase in Barley, Wheat, and Rye and Their Organization in the Barley Genome

The HemA genes encoding glutamyl-tRNA reductase, an important enzyme in chlorophyll and heme biosynthesis, are located on barley chromosome 5 (1H), homoeologous chromosomes 1A, IB, and ID of common wheat, and rye chromosome 1R. Southern blot analyses revealed a low copy multigene family for HemA genes in the barley genome.

A PCR-based marker for targeting small rye segments in wheat background

We attempted to develop a PCR-based marker that detects various segments of rye chromosome incorporated into wheat. We designed three sets of PCR primers based on the nucleotide sequence data of a rye repetitive sequence previously reported. One of the primer sets amplified a clear ca. 1.4 kb fragment in a rye cultivar but not in any form of wheat, diploid, tetraploid or hexaploid. We used this critical primer set for PCR of various wild species and cultivars of rye, an array of wheat plants carrying different rye chromosomes or small segments from different regions of rye chromosome 1R, and plants carrying parts of the rye B chromosome. The PCR amplified the 1.4 kb fragment in all the plant materials examined. We believe this PCR primer set will be useful as a universal PCR-based marker for the introgression of rye chromosome segments in the wheat genome.

Substitution analysis of seedling stage copper tolerance in wheat

The relatively copper-tolerant wheat variety Chinese Spring (recipient), the copper-sensitive variety Cappelle Desprez (donor) and their substitution lines were screened for copper tolerance in a soil pot experiment under artificial growth conditions. Chromosomes 5A, 5B, 5D and 7D of Cappelle Desprez significantly decreased the copper tolerance of the recipient variety to varying extents. By contrast, the 6B and 3D chromosomes significantly increased the copper tolerance of Chinese Spring, suggesting that a wide range of allelic differences could be expected between wheat genotypes for this character. The significant role of homologous group 5 in copper tolerance was confirmed by testing wheat-rye substitution lines. The substitution of rye chromosome 5R (5R/5A substitution line) into a wheat genetic background significantly increased the copper tolerance of the recipient wheat genotype. The results suggest that chromosomes 5R and 5A probably carry major genes or gene complexes responsible for copper tolerance, and that the copper tolerance of wheat can be improved through the substitution of a single chromosome carrying the responsible genes. At the same time, it is also possible that the effect of homologous group 5 is not specific to copper tolerance, but that the genes located on these chromosomes belong to a general stress adaptation (frost, cold, vernalisation requirements, etc.) complex, which has already been detected on this chromosome. To answer this question further studies are needed to determine the real effect of these chromosome regions and loci on copper tolerance.

Analysis and sorting of rye (Secale cerealeL.) chromosomes using flow cytometry

Procedures for chromosome analysis and sorting using flow cytometry (flow cytogenetics) were developed for rye (Secale cereale L.). Suspensions of intact chromosomes were prepared by mechanical homogenization of synchronized root tips after mild fixation with formaldehyde. Histograms of relative fluorescence intensity obtained after the analysis of DAPI-stained chromosomes (flow karyotypes) were characterized and the chromosome content of the DNA peaks was determined. Chromosome 1R could be discriminated on a flow karyotype of S. cereale 'Imperial'. The remaining rye chromosomes (2R-7R) could be discriminated and sorted from individual wheat-rye addition lines. The analysis of lines with reconstructed karyotypes demonstrated a possibility of sorting translocation chromosomes. Supernumerary B chromosomes could be sorted from an experimental rye population and from S. cereale 'Adams'. Flow-sorted chromosomes were identified by fluorescence in situ hybridization (FISH) with probes for various DNA repeats. Large numbers of chromosomes of a single type sorted onto microscopic slides facilitated detection of rarely occurring chromosome variants by FISH with specific probes. PCR with chromosome-specific primers confirmed the identity of sorted fractions and indicated suitability of sorted chromosomes for physical mapping. The possibility to sort large numbers of chromosomes opens a way for the construction of large-insert chromosome-specific DNA libraries in rye.

Genetic mapping of Dn7, a rye gene conferring resistance to the Russian wheat aphid in wheat.

The Russian wheat aphid is a significant pest problem in wheat and barley in North America. Genetic resistance in wheat is the most effective and economical means to control the damage caused by the aphid. Dn7 is a rye gene located on chromosome 1RS that confers resistance to the Russian wheat aphid. The gene was previously transferred from rye into a wheat background via a 1RS/1BL translocation. This study was conducted to genetically map Dn7 and to characterize the type of resistance the gene confers. The resistant line '94M370' was crossed with a susceptible wheat cultivar that also contains a pair of 1RS/1BL translocation chromosomes. The F(2) progeny from this cross segregated for resistance in a ratio of 3 resistant: 1 susceptible, indicating a single dominant gene. One-hundred and eleven RFLP markers previously mapped on wheat chromosomes 1A, 1B and 1D, barley chromosome 1H and rye chromosome 1R, were used to screen the parents for polymorphism. A genetic map containing six markers linked to Dn7, encompassing 28.2 cM, was constructed. The markers flanking Dn7 were Xbcd1434 and XksuD14, which mapped 1.4 cM and 7.4 cM from Dn7, respectively. Dn7 confers antixenosis, and provides a higher level of resistance than that provided by Dn4. The applications of Dn7 and the linked markers in wheat breeding are discussed.

Comparative effects of rye chromosomes 1R and 5R on androgenesis in cultured anthers of wheat-rye substitution lines dependending on the line origin

The effects of rye chromosomes 1R and 5R on androgenesis in cultured anthers of wheat--rye substitution lines was studied as dependent on the cultivar origin of the rye chromosomes and on the wheat genome (A or D) subjected to substitution. Chromosome 1R stimulated embryogenesis in anther cultures, while chromosome 5R suppressed it regardless of whether the corresponding wheat chromosomes were substituted in the A or D genome. The effect of chromosome 1R on embryogenesis proved to depend on its cultivar origin. Along with rye chromosome 1R, wheat chromosome 1A was shown to substantially affect total seedling regeneration. Regeneration of green seedlings was dramatically affected both by rye chromosome 1R and by wheat chromosome 1D. The results supported the published data that individual androgenesis parameters (embryogenesis, total plant regeneration, green plant regeneration) are controlled by different genetic mechanisms.