Chromosome pairing in durum wheat haploids with and without <Emphasis Type="Italic">ph1b</Emphasis> of bread wheat

Durum or macaroni wheat (Triticum turgidum L., 2n = 4x = 28; AABB) is an allotetraploid with two related genomes, AA and BB, each with seven pairs of homologous chromosomes. Although the corresponding chromosomes of the two genomes are potentially capable of pairing with one another, the Ph1 (Pairing homoeologous) gene in the long arm of chromosome 5B permits pairing only between homologous partners. As a result of this Ph1-exercised disciplinary control, durum wheat and its successor, bread wheat (Triticum aestivum L., 2n = 6x = 42; AABBDD) show diploid-like chromosome pairing and hence disomic inheritance. The Ph mutants in the form of deletions are available in bread wheat (ph1b) and durum wheat (ph1c). Thus, ph1b-haploids of bread wheat and ph1c-haploids of durum wheat show extensive homoeologous pairing that has been shown by us and several others. Here we study the effect of ph1b allele of bread wheat on chromosome pairing in durum haploids, whereas we studied earlier the effect of ph1c allele in durum haploids that we synthesized. In durum wheat, the ph1b-haploids show much higher (49.4% of complement) pairing than the ph1c-haploids (38.6% of complement). Mention of a trademark or proprietary product does not constitute a guarantee or warranty of the product by the USDA or imply approval to the exclusion of other products that also may be suitable.     

Characterization of wheat-alien translocations conferring resistance to diseases and pests: current status

Summary Wild relatives of common wheat, Triticum aestivum, and related species are an important source of disease and pest resistance and several useful traits have been transferred from these species to wheat. C-banding and in situ hybridization analyses are powerful cytological techniques allowing the detection of alien chromatin in wheat. C-banding permits identification of the wheat and alien chromosomes involved in wheat-alien translocations, whereas genomic in situ hybridization analysis allows determination of their size and breakpoint positions. The present review summarizes the available data on wheat-alien transfers conferring resistance to diseases and pests. Ten of the 57 spontaneous and induced wheat-alien translocations were identified as whole arm translocations with the breakpoints within the centromeric regions. The majority of transfers (45) were identified as terminal translocations with distal alien segments translocated to wheat chromosome arms. Only two intercalary wheat-alien transloctions were identified, one induced by radiation treatment with a small segment of rye chromosome 6RL (H25) inserted into the long arm of wheat chromosome 4A, and the other probably induced by homoeologous recombination with a segment derived from the long arm of a group 7 Agropyron elongatum chromosome with Lr19 inserted into the long arm of 7D. The presented information should be useful for further directed chromosome engineering aimed at producing superior germplasm.Contribution No. 96-55-J from the Kansas Experimental Station, Kansas State University, Manhattan, KS 66506-5502, USA.     

Control and inheritance of resistance to yellow rust in Triticum aestivum-Lophopyrum elongatum chromosome substitution lines

A set of T. aestivum-L. elongatum chromosome substitution lines was tested for yellow rust resistance at the seedling stage. Inheritance of the resistance and esterase-5 (Est-5) variation were studied. The results demonstrated that L.elongatum carried a new gene(s) conferring yellow rust resistance. This gene was dominant and located on chromosome 3E of L. elongatum. The biochemical locus encoding Est-5was also located on chromosome 3E, and co-segregated with theYr gene(s) in the wheat background. The transmission frequencies of chromosome 3E in 3E(3A) × CS, 3E(3B) × CS and 3E(3D) × CS hybrids were scored.None of the hybrids transmitted the alien chromosome at thetheoretical maximum rate, but the transmission frequencies ofchromosome 3E in F₂ populations of 3E(3A) × CS and 3E(3D) × CS were significantly higher than in thatof 3E(3B) × CS.     

Transfer of rust resistance from Aegilops ovata into bread wheat (Triticum aestivum L.) and molecular characterisation of resistant derivatives

An interspecific cross was made to transfer leaf rust and stripe rust resistance from an accession of Aegilops ovata (UUMM) to susceptible Triticum aestivum (AABBDD) cv. WL711. The F₁was backcrossed to the recurrent wheat parent, and after two to three backcrosses and selfing, rust resistant progenies were selected. The C-banding study in a uniformly leaf rust and stripe rust resistant derivative showed a substitution of the 5M chromosome of Ae. ovata for 5D of wheat. Analysis of rust resistant derivatives with mapped wheat microsatellite makers confirmed the substitution of 5M for 5D. Some of these derivatives also possessed one or more of the three alien translocations involving 1BL, 2AL and 5BS wheat chromosomes which could not be detected through C-banding. A translocation involving 5DSof wheat and the substituted chromosome 5M of Ae. ovata was also observed in one of the derivatives. Susceptibility of this derivative to leaf rust showed that the leaf rust resistance gene(s) is/are located on short arm of 5M chromosome of Ae. ovata. Though the Ae. ovatasegment translocated to 1BL and 2AL did not seem to possess any rust resistance gene, the alien segment translocated to 5BS may also possess gene(s) for rust resistance. The study demonstrated the usefulness of microsatellite markers in characterisation of interspecific derivatives. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic analysis of the Aegilops longissima 3S chromosome carrying the Pm13 resistance gene

The distal region of the short arm of chromosome 3S from Aegilopslongissima, which carries the powdery mildew resistance gene Pm13, was introgressed into common wheat. Due to suppression of recombination between this region and corresponding wheat homoeologous segments, a possible strategy to construct a genetic map around the Pm13 gene was based on crosses between a wheat addition line carrying the Ae.longissima 3S chromosome and the corresponding 3S addition lines of Ae.searsii and Ae. variabilis. The efficiency of this strategy was evaluated by scoring recombination frequencies inprogenies derived from these crosses. Recombination between 3S chromosomes fromAe. searsii and Ae. longissimawas very low, whereas 26.5% recombinant alien chromosomes were obtained from the cross involving the Ae. variabilisand Ae. longissima 3S addition lines. These data were used to construct a3S chromosome map that resulted largely colinear to the consensus map of the homoeologous group 3 of wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Molecular mapping and detection of the yellow rust resistance gene Yr26 in wheat transferred from Triticum turgidum L. using microsatellite markers

Yellow rust (stripe rust), caused by Puccinia striiformis Westend f. sp. tritici, is one of the most devastating diseases of wheat throughout the world. Wheat-Haynaldia villosa 6AL.6VS translocation lines R43, R55, R64 and R77, derived from the cross of three species, carry resistance to both yellow rust and powdery mildew. An F₂ population was established by crossing R55 with the susceptible cultivar Yumai 18. The yellow rust resistance in R55 was controlled by a single dominant gene, which segregated independently of the powdery mildew resistance gene Pm21 located in the chromosome 6VS segment, indicating that the yellow rust resistance gene and Pm21 are unlikely to be carried by the same alien segment. This yellow rust resistance gene was considered to beYr26, originally thought to be also located in chromosome arm 6VS. Bulked Segregation Analysis and microsatellite primer screens of the population F₂ of Yumai 18 × R55 identified three chromosome 1B microsatellite locus markers, Xgwm11, Xgwm18 and Xgwm413, closely linked to Yr26. Yr26 was placed 1.9 cM distal of Xgwm11/Xgwml8, which in turn were 3.2 cM from Xgwm413. The respective LOD values were 21 and 36.5. Therefore, Yr26 was located in the short arm of chromosome 1B. The origin and distribution of Yr26 was investigated by pedigree, inheritance of resistance and molecular marker analysis. The results indicated that Yr26 came from Triticum turgidum L. Three other 6AL.6VS translocation lines, R43, R64 and R77, also carried Yr26. These PCR-based microsatellite markers were shown to be very effective for the detection of the Yr26 gene in segregating populations and therefore can be applied in wheat breeding. This revised version was published online in July 2006 with corrections to the Cover Date.     

Transfer of the Glu-D1 Gene from Chromosome 1D to Chromosome 1A in Hexaploid Triticale

To complement previously developed recombinant chromosomes 1R.1D, two series of translocations involving the Glu-D1 gene from chromosome ID to chromosome 1A were produced in hexaploid triticale. These series involve seven independent transfers of allele d encoding for high molecular weight glutenin subunits 5+10 and ten independent transfers involving allele a encoding for HMW glutenin subunits 2 + 12. The frequency of homoeologous recombination between chromosomes 1A and 1D was within the range observed for pairs of homologues in wheat, supporting earlier observations that homoeologous recombination in triticale is frequent. Recombined chromosomes 1A.1D can be used to introduce the Glu-D1 gene to durum wheats, and to manipulate the dosage of Glu-D1 in hexaploid triticale and bread wheat.     

Chromosome location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L.) 1. Mlk and other alleles at the Pm3 locus

Summary Several wheat cultivars/lines were inoculated with isolates of Erysiphe graminis tritici to identify new genes/alleles for resistance. The wheats were tested with 13 isolates that had been characterized from responses on differential lines with known resistance genes. Gene Mlk which occurs in cultivars Kolibri, Syros, Ralle and several other European common wheats was found to be an allele at the Pm3 locus and is now designated Pm3d. The mildew resistance in an old Australian wheat, W150, is conferred by a single gene also allelic to Pm3 and now designated Pm3e. The near-isogenic line Michigan Amber/8^*Cc possesses another allele now designated Pm3f. A Syrian land variety of common wheat shows mildew resistance that is conditioned by the combination of genes Pm1 and Pm3a. Finally, two accessions of Triticum aestivum ssp. sphaerococcum appeared to possess the Pm3c allele.     

The introduction into bread wheat of a major gene for resistance to powdery mildew from wild emmer wheat

Summary A new source of resistance to wheat powdery mildew caused by Erysiphe graminis has been transferred to hexaploid bread wheat, Triticum aestivum, from the wild tetraploid wheat, Triticum dicoccoides. The donor was crossed to bread wheat and the pentaploid progeny was then self-pollinated. Plants having a near stable hexaploid chromosome complement were selected in the F₃ progeny and topcrossing and backcrossing of these to a second wheat cultivar to improve the phenotype was undertaken. Monosomic analysis of early backcross lines showed the transferred gene to be located on chromosome 4A. The gene has been designated Pm16.     

Chromosomal location of genes for resistance to powdery mildew in common wheat (Triticum aestivum L. em Thell.) 7. Gene Pm29 in line Pova

Chromosomal localization and linkage mapping of a powdery mildewresistance gene were conducted in the resistant wheat line Pova, derivedfrom a Triticum aestivum cv. Poros-Aegilops ovata-alien additionline. Monosomic analysis revealed that a major dominant gene was locatedon chromosome 7D. This gene possessed a distinct disease response patternagainst a differential set of Blumeria graminis tritici isolates andsegregated independently from resistance gene Pm19 also located onwheat chromosome 7D. Molecular genetic analysis showed that theresistance gene in Pova was specifically located on the long arm ofchromosome 7D closely linked to one RFLP and three AFLP markers. It isproposed that the new gene be designated Pm29.     

Sources of resistance to Cephalosporium gramineum in Triticum and Agropyron species

Summary Resistance to the soil-borne pathogen Cephalosporium gramineum was evaluated in Agropyron elongatum, A. intermedium. A. intermedium var. trichophorum, an Agrotriticum, and eight species of Triticum. Only A. elongatum and A. intermedium showed high levels of resistance. Agrotriticum (56 chromosomes) was resistant too. High resistance to C. gramineum is available, but its utilization will probably require the use of chromosome substitution techniques to transfer the resistance into an agronomically useful wheat.     

Induction of a mutation in the male fertility gene of the preferentially transmitted Aegilops sharonensis chromosome 4S1 and its application for hybrid wheat production

Summary Wheat plants nullisomic for chromosome 4B are male sterile due to the absence of the male fertility gene Ms1. However, plants in which chromosome 4B has been substituted by the preferentially transmitted chromosome 4S¹ of Ae. sharonensis are male fertile due to the compensating effect of Ms4 on the alien chromosome. This substitution line has been mutated and three recessive mutation of Ms4 have been selected. Plants homozygous for these mutations are male sterile. The implication of these mutations for hybrid wheat production is discussed.     

Identification and genetic mapping of a powdery mildew resistance gene in wild emmer (<Emphasis Type="Italic">Triticum dicoccoides</Emphasis>) accession IW72 from Israel

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is a devastating disease of wheat (Triticum aestivum) in China and worldwide, causing severe yield losses annually. Wild emmer (T. dicoccoides) accession IW72 collected from Israel is resistant to powdery mildew at the seedling and adult stages. Genetic analysis indicated that the resistance was controlled by a single dominant gene, temporarily designated MlIW72. The F₂ population and F₃ families derived from a hybrid between IW72 and susceptible durum wheat line Mo75 were used for molecular mapping of the resistance gene. MlIW72 was linked with SSR loci Xgwm344, Xcfa2040, Xcfa2240, Xcfa2257 and Xwmc525 on the long arm of chromosome 7A. In addition, two STS markers, MAG2185 (derived from RFLP marker PSR680) and MAG1759 (developed from EST CD452874), were mapped close to MlIW72. All these markers were physically located in the terminal bin 0.86–1.00 of 7AL. The chromosome location and genetic mapping results suggested that the powdery mildew resistance gene identified in wild emmer accession IW72 might be a new allele at the Pm1 locus or a new locus closely linked to Pm1.     

Sources of resistance to Mayetiola Destructor in breadwheat and durum wheat in Morocco

Summary Grain yield reductions of both breadwheat (Triticum aestivum L.) and durum wheat (Triticum turgidum L. var. durum) caused by attacks of Hessian fly (Mayetiola destructor Say) are second perhaps only to those caused by inadequate soil moisture in Morocco. To identify effective sources of resistance, 817 entries of common wheat and durum wheat reported to be resistant to Hessian fly were evaluated under natural infestations in Morocco. A large number of genes conferring virulence are present in populations of Moroccan Mayetiola. The genes H₁, H₂, H₃, h₄, H₆, H₇, H₈, H₉, H₁₀, H₁₁, H₁₄, H₁₅, and H₁₆ as well as the Marquillo, Kawvale and PI 94587 resistance sources are not useful for cereal improvement in North Africa. Luso, which has the gene H₁₂, also appeared susceptible in limited testing. Genotypes having the genes H₅ and H₁₃ were identified as significantly reducing larval survial in natural populations of Mayetiola. Of 11 resistant breadwheats identified with unknown genes, seven were from Portugal and three were from the Soviet Union. Although none of the durums tested had high levels of reistance, the two most promising durums were from Portugal. It is proposed that initially H₅ be deployed in durum wheats and H₁₃ be used in common wheat improvement. Leaf pubescence appears of little use in reducing the larval survival of Mayetiola.     

Effect of alien 5R(5A) chromosome substitution on ear-emergence time and winter hardiness in wheat-rye substitution lines

Ear emergence time and response to vernalization were investigated in 12 alien substitution lines in which a pair of chromosomes 5A of recipient spring wheat cultivars was replaced by a pair of chromosomes 5R of Siberian spring rye ‘Onokhoiskaya’. The recipients were 12 spring cultivars of common wheat, each carrying different Vrn genes. Spring rye ‘Onokhoiskaya’ had the Sp1 (now called Vrn-R1) gene for spring growth habit located on chromosome 5R, but its expression was weaker. The Vrn-R1 gene had no effect on growth habit, ear emergence time and response to vernalization in wheat-rye substitution lines. Ears emerged significantly later in the 5R(5A) alien substitution lines than in the recipient wheat cultivars with the Vrn-A1/Vrn-B1/vrn-D1 or Vrn-A1/vrn-B1/Vrn-D1 genotypes. No difference in ear emergence time was found between most of the 5R(5A) alien substitution lines and the cultivars carrying the recessive vrn-A1 gene. The presence of the Vrn2a and Vrn2b alleles at the Vrn2 (now called Vrn-B1) locus located on wheat chromosome 5B was confirmed.The replacement of chromosome 5A by chromosome 5R in wheat cultivars ‘Rang’ and ‘Mironovskaya Krupnozernaya’, which carries the single dominant gene Vrn-A1, converted them to winter growth habit. In field studies near Novosibirsk the winter hardiness of 5R(5A) wheat–rye substitution lines of ‘Rang’ and ‘Mironovskaya Krupnozernaya’ was increased by 20–47% and 27–34%, respectively, over the recurrent parents.     

Ph I-induced transfer of leaf and stripe rust-resistance genes from Aegilops triuncialis and Ae. geniculata to bread wheat

Leaf and stripe rusts are severe foliar diseases of bread wheat. Recently, chromosomes 5M^g from the related species Aegilops geniculata that confers resistance to both leaf and stripe rust and 5U^t from Ae. triuncialis conferring resistance to leaf rust have been transferred to bread wheat in the form of disomic DS5M^g(5D) and DS5U^t(5A) chromosome substitution lines. The objective of this study was to shorten the alien segments in these lines using Ph ^I-mediated, induced homoeologous recombination. Putativerecombinants were evaluated for their rust resistance, and by genomic in situ hybridization and microsatellite analyses. One agronomically useful wheat-Ae. geniculata recombinant resistant to leaf and stripe rust was identified that had only a small terminal segment of the 5M^gL arm transferred to the long arm of an unidentified wheat chromosome. This germplasm can be used directly in breeding programs. Only one leaf rust-resistant wheat-Ae. triuncialis recombinant, which consists of most of the complete 5U^t chromosome with a small terminal segment derived from 5AS, was identified. This germplasm will need further chromosome engineering before it can be used in wheat improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Karyological characterization of a partial amphiploid,Triticum turgidum L. var. durum × Agropyron intermedium (Host) P.B.

Summary A Giemsa-C-banded karyotype of a partial amphiploid, Triticum turgidum L. var. durum cv. Nodak × Agropyron intermedium (Host) P.B., called MT-2, was analyzed. MT-2 is a winterhardy grasslike octoploid germplasm which survived 5 winters in Montana, and its seed weight is 3 times that of A. intermedium seed. The MT-2 C-banding karyotype shows 6 chromosome pairs each of the A and B wheat genomes with 3A and 4B missing. Chromosomes 1B and 2B are involved in a reciprocal homozygous translocation (T1BS·2BS, T1BL·2BL) which was also confirmed by a nucleolus-associated quadrivalent in an MT-2 × durum wheat backcross. In addition to the wheat chromosomes, MT-2 consistently shows 16 A. intermedium chromosome pairs which are designated from A to P. These chromosomes show C-banding patterns similar to those reported earlier in the literature. A large amount of C-banding polymorphism and structural rearrangements in A. intermedium itself presently make a definite chromosome assignment to the homeologous groups of the Triticeae difficult. The data presented are crucial for further directed manipulation of this germplasm aimed at producing valuable chromosome additions and substitutions in wheat.contribution No. J-2767 from Montana Agric. Exp. Stn.     

Fluorescent in situ hybridization — a useful aid to the introduction of alien genetic variation into wheat

Summary Fluorescent in situ hybridization (FISH) of DNA to plant chromosomes has proved to be a powerful cytogenetic tool. The value of fluorescent in situ hybridization of total genomic DNA (GISH) of related species is demonstrated in the determination of wheat/alien chromosome pairing in hybrids. Its use for assessing the relative merits of the various genes that affect chromosome pairing is also shown.The ability of GISH to identify the presence in wheat of whole alien chromosomes or alien chromosome segments is illustrated. The potential of FISH for detecting repeated DNA sequences, low copy sequences and single copy genes is discussed.Abbreviations FISH fluorescent in situ hybridization - GISH genomic in situ hybridization - PRINS primer-induced in situ hybridization     

Hybrids between durum wheat and Thinopyrum junceiforme: Prospects for breeding for scab resistance

Tetraploid wheatgrass, Thinopyrum junceiforme(2n = 4x = 28; J₁J₁J₂J₂), a wild relative of wheat, is an excellent source of resistance to Fusarium head blight. Intergeneric F₁ hybrids (2n = 4x = 28; ABJ₁J₂) between durum wheat (Triticum turgidum; 2n = 4x = 28; AABB) cultivars Lloyd or Langdon and Th. junceiforme were synthesized. Most of the pairing in F₁ hybrids was between the J₁- and J₂-genome chromosomes. Some pairing occurred between wheat chromosomes and alien chromosomes, resulting in segmental exchange that was confirmed by fluorescent in situ hybridization (FISH). The F₁hybrids were largely male-sterile and were backcrossed, as the female parent, to the respective durum cultivar. Backcrosses from Lloyd × Th. junceiforme hybrids yielded fertile partial amphiploids (2n = 6x = 42; AABBJ₁J₂) as a result of functioning of unreduced female gametes of the hybrid. Lloyd proved to be a more useful durum parent than Langdon in crosses with Th. junceiforme designed to transfer scab resistance genes. Pairing in the amphiploids was characterized by preferential pairing,which resulted in bivalent formation. However, some intergeneric pairing also occurred. Several fertile hybrid derivatives were produced by further backcrossing and selfing. The introduction of alien chromatin into the durum complement was confirmed by FISH. Hybrid derivative lines had significantly lower mean infection scores (p = 0.01), the best showing 10.93% infection, whereas the parental durum cultivars had 70.34% to 89.46% infection. Hybridization with wild relatives may offer an excellent means of introducing scab resistance into durum wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Cytological analyses of hybrids and derivatives of hybrids between durum wheat and Thinopyrum bessarabicum,using multicolour fluorescent GISH*

The wheat progenitors and other wild relatives continue to be important sources of genes for agronomically desirable traits, which can be transferred into durum wheat (Triticum turgidum; 2n = 4x = 28; AABB genomes) cultivars via hybridization. Chromosome pairing in durum × alien species hybrids provides an understanding of genomic relationships, which is useful in planning alien gene introgression strategies. Two durum cultivars, ‘Lloyd’ and ‘Langdon’, were crossed with diploid wheatgrass, Thinopyrum bessarabicum (2n = 2x = 14; JJ), to synthesize F₁ hybrids (2n = 3x = 21; ABJ) with Ph1. ‘Langdon’ disomic substitution 5D(5B) was used as a female parent to produce F₁ hybrids without Ph1, which resulted in elevation of pairing between durum and grass chromosomes – an important feature from the breeding standpoint. The F₁ hybrids were backcrossed to respective parental cultivars and BC₁ progenies were raised. ‘Langdon’ 5D(5B) substitution × Th. bessarabicum F₁ hybrids were crossed with normal ‘Langdon’ to obtain BC₁ progeny. Chromosome pairing relationships were studied in F₁ hybrids and BC₁ progenies using both conventional staining and fluorescent genomic in situ hybridization (fl‐GISH) techniques. Multicolour fl‐GISH was standardized for characterizing the nature and specificity of chromosome pairing: A–B, A–J and B–J pairing. The A–J and B–J pairing will facilitate gene introgression in durum wheat. Multicolour fl‐GISH will help in characterizing alien chromosome segments captured in the durum complement and in their location in the A and/or B genome, thereby accelerating chromosome engineering research.