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.     

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.     

Leaf rust and stripe rust resistance genes Lr54 and Yr37 transferred to wheat from Aegilops kotschyi

The tendency of unpaired meiotic chromosomes to undergo centric misdivision was exploited to translocate leaf rust and stripe rust resistance genes from an Aegilops kotschyi addition chromosome to a group 2 chromosome of wheat. Monosomic and telosomic analyses showed that the translocation occurred to wheat chromosome arm 2DL. The introgressed region did not pair with the corresponding wheat 2DL telosome during meiosis suggesting that a whole arm may have been transferred. Female transmission of the resistance was about 55% whereas male transmission was strongly preferential (96%). The symbols Lr54 and Yr37 are proposed to designate the new resistance genes.     

Transfer of disease resistance genes from Triticum araraticum to common wheat

The wild tetraploid wheat species Tr$$ (Zhuk) Zhuk Var. araratieum is a source of pest resistance genes for T$$ aesti$$ L. Our objectives were to describe the breeding behaviour of T.arartuititm when backcrossed to common wheat and transfer resistance to leaf rust (caused by Pu$$) and powdery mildew (caused by Blumeria $$wheat. Crosses were made between five wheat genotypes and $$ accessions. Fertifity and chromosome numbers of BC$$_; plants were determined. Resistance to leaf rust was transferred toBC₂ -derived families from 10 different T’ararati$$an accessions. Leaf rust resistance genes in nine T. araratieum accessions can be assigned to at least four loci. Leaf rust resistance transferred from three accessions was inherited in the hexaploid derivatives as a single. $$ gene in each case. Resistance to powdery mildew was also detected in the T. araratie$$ backcross derivatives. Fertile hexaploid derivatives expressing T’araratieum-derived resistance genes can be recovered after two backcrosses to wheat cultivars.     

Detection of molecular markers linked to the durable adult plant stripe rust resistance gene Yr18 in bread wheat (Triticum aestivum L.)

Stripe rust of wheat caused by Puccinia striiformis West. f. sp. tritici presents a serious problem for wheat production worldwide, and identification and deployment of resistance sources to it are key objectives for many wheat breeders. Here we report the detection of simple sequence repeat (SSR) markers linked to the durable adult plant resistance of cv. ‘Otane’, which has conferred this resistance since its release in New Zealand in 1984. A double haploid population from a cross between ‘Otane’ and the susceptible cv. Tiritea’ was visually assessed for adult plant infection types (IT) in the glasshouse and field, and for final disease severity in the field against stripe rust pathotype 106E139A⁺. At least three resistance loci controlled adult plant resistance to stripe rust in this population. Quantitative trait loci (QTL) mapping results revealed that two of these, one on chromosome 7DS corresponds to the durable adult plant resistance gene Yr18 and other on chromosome 5DL were contributed from ‘Otane’; while the remaining one on chromosome 7BL, was contributed from the susceptible ‘Tiritea’. Interval mapping placed the ‘Otane’‐resistant segment near the centromere of chromosome 7DS at a distance of 7 cM from the SSR marker gwm44. The stability of QTL in the two environments is discussed. SSR gwm44 is potentially a candidate marker for identifying the durable resistance gene Yr18 in breeding programmes.     

The Transfer of Thinopyrum-Derived Leaf-rust Resistance from Common Wheat to Durum Wheat.

The leaf rust resistance gene on chromosome 7AL of ‘Chinese Spring’ transfer no. 12 derived from Thinopyrum ponticum, was transferred to durum wheat by standard backcrossing. In ‘Agatha’ and ‘Indis’ a leaf rust resistance gene from Thinopyrum ponticum and Thinopyrum ponticum respectively, is found on a translocated segment on chromosome arm 7DL. The use of the ‘Langdon’ disomic D-chromosome substitution lines for 7A and 7B resulted in the recovery of tetraploid leaf-rust resistant lines from the crosses with ‘Agatha’ in the B₂F₁ generation. Tetraploid lines carrying the ‘Indis’ translocation segment were recovered in the B₂F₂ generation. The F₂ segregation ratios for rust resistance after selfing or back-crossing generally fitted a 1: 1 ratio indicating non-transmission of the translocation segments in the male gametes. Homozygous resistant plants were not obtained. Meiotic instability was observed in 28 chromosome B₂ F₂ derivatives of the crosses between ‘Chinese Spring’ transfer no. 12 and durum wheat.     

Monosomic analyses of stripe rust and leaf rust resistance genes in winter wheat varieties Lüquyu and Yantar

Summary A set of 21 monosomics of Novosadska Rana-1 was used to locate the rust resistance genes of Lüqiyu, a stripe rust resistant line developed by BAU and Yantar, a leaf rust resistant wheat introduced from Bulgaria. The resistance of the former to p. striiformis race C25 was conditioned by a dominant gene located on chromosome 2B, whereas that of the latter to P. recondita race CL3 was controlled by two complementary dominant genes located on chromosomes 5A and 1D, respectively. The relationship of the stripe rust resistance gene in Lüqiyu to Yr5, Yr7 or Yr _Suwon' all located on chromosome 2B is unknown. The two complementary leaf rust resistance factors in Yantar appear to be new.     

Inheritance and chromosome location of leaf rust resistance in durum wheat cultivar Wollaroi

The Australian durum cultivar Wollaroi produced low leaf rust responses in the field since its release in 1993. A recombinant inbred line population was developed from a cross of Wollaroi with a susceptible landrace Bansi. Monogenic inheritance of low seedling leaf rust response against the Puccinia triticina pathotype 104-1,2,3,6,(7),(11),13 was observed. This gene was temporarily designated as LrWo. A DArT based map of Wollaroi/Bansi was used to determine the genomic location of LrWo and it was mapped in chromosome 5BS. Following enrichment of the DArT map of chromosome 5BS with SSR markers, LrWo was flanked by gwm234 (7.2 cM) and wPt-1420 (20.3 cM) distally and proximally, respectively. A previously characterised gene Lr52 was also located on the chromosome arm 5BS, proximal to gwm234. Based on genetic association with the marker gwm234 at a similar distance, we concluded that LrWo could be either Lr52 or is another allele of this locus. Based on infection type comparison the latter argument seems more plausible.     

Development of a PCR assay and marker-assisted transfer of leaf rust resistance gene <Emphasis Type="Italic">Lr58</Emphasis> into adapted winter wheats

Leaf rust resistance gene Lr58 derived from Aegilops triuncialis L. was transferred to the hard red winter wheat (HRWW) cultivars Jagger and Overley by standard backcrossing and marker-assisted selection (MAS). A co-dominant PCR-based sequence tagged site (STS) marker was developed based on the sequence information of the RFLP marker (XksuH16) diagnostically detecting the alien segment in T2BS·2BL-2^tL(0.95). STS marker Xncw-Lr58-1 was used to select backcross F₁ plants with rust resistance. The co-dominant marker polymorphism detected by primer pair NCW-Lr58-1 efficiently identified the homozygous BC₃F₂ plants with rust resistance gene Lr58. The STS marker Xncw-Lr58-1 showed consistent diagnostic polymorphism between the resistant source and the wheat cultivars selected by the US Wheat Coordinated Agricultural Project. The utility and compatibility of the STS marker in MAS programs involving robust genotyping platforms was demonstrated in both agarose-based and capillary-based platforms. Screening backcross derivatives carrying Lr58 with various rust races at seedling stage suggested the transferred rust resistance in adapted winter wheats is stable in both cultivar backgrounds. Lr58 in adapted winter wheat backgrounds could be used in combination with other resistance genes in wheat rust resistance breeding.     

Attempts to remove gametocidal genes co-transferred to common wheat with rust resistance from <Emphasis Type="Italic">Aegilops speltoides</Emphasis>

Rust resistance genes (introgressions S24 and S13) transferred to hexaploid wheat from two Aegilops speltoides accessions could not be used commercially due to associated gametocidal (Gc) genes. Crosses to wheat followed by rigorous selection for increased fertility were employed in an attempt to separate the unmapped S24 stem rust resistance from the Gc gene(s). However, improved fertility of the better selections could not be maintained in subsequent generations. Since the S13 introgression (leaf, stripe and stem rust resistances) mapped to chromosome 3A, allosyndetic pairing induction was used in an attempt to remove the Gc gene(s). This produced putative primary recombinants with improved fertility and plant type, the best of which had exchanged a small region of Ae. speltoides chromatin, yet was still associated with (reduced) Gc effects. This selection (04M127-3, which appears to have the Su1-Ph1 suppressor) was then crossed with wheat. Surprisingly, the 04M127-3 gametocidal effect differed drastically from that of the original introgression allowing the recovery of 35 recombinant, leaf rust resistant progeny. Microsatellite and DArT markers showed that each secondary recombinant had exchanged most of the Ae. speltoides chromatin. Although the data suggested that a complex multigenic interaction may govern the gametocidal response, preliminary indications are that the Gc effect had largely been removed and it now seems possible to completely separate the gametocidal genes from the S13 leaf rust resistance gene (here designated Lr66). The associated (S13) stripe rust and stem rust resistance genes were lost during recombination.     

Alleles on the two dwarfing loci on 4B and 4D are main drivers of FHB‐related traits in the Canadian winter wheat population “Vienna” × “25R47”

Fusarium head blight (FHB), leaf rust and stem rust are among the most destructive wheat diseases. High‐yielding, native disease resistance sources are available in North America. The objective of this study was to map loci associated with FHB traits, leaf rust, stem rust and plant height in a “Vienna”/”25R47” population. DArT markers were used to generate a genetic map, and quantitative trait loci (QTL) analysis was performed by evaluating 113 doubled haploid lines across three environments in Ontario, Canada. FHB resistance QTL were identified on chromosomes 4D, 4B, 2D and 7A, while a QTL for leaf and stem rust resistance was identified on chromosome 1B. The dwarfing alleles of both Rht‐B1 and Rht‐D1 were associated with increased FHB index and DON content.     

Molecular and agronomic characterization of wheat-Agropyron intermedium recombinant chromosomes

Thirty‐six wheat‐Agropyron intermedium (host) Beauv. [Syn. Trichopyrum intermedium (host) A. Love, Elytrigia intermedia (host) Nevski, Thinopyrum intermedium (host) Barkworth and Dewey] 7A/7Ai‐1 recombinant chromosomes were characterized using DNA markers. Analysis of recombinant chromosomes using 15 restriction fragment length polymorphism probes identified the homoeologous crossover products that had varying length of A. intermedium chromatin introgressed onto chromosome 7A of common wheat. The linear order of the probe loci was established along the lengths of the chromosomes. The short arm recombinants that had A. intermedium chromatin distal to the locus Xpsr108 and proximal to the locus Xpsr119 were resistant to wheat stem rust, indicating that the rust resistance gene (Sr44) was located on the distal part of chromosome arm 4Ai‐1s. The barley yellow dwarf virus (BYDV) resistance gene reported to be present on the long arm of chromosome 7Ai‐1 was found to be ineffective against the BYDV serotype used in the present study.     

Identification of recombinants carrying stripe rust resistance gene Yr57 and adult plant stem rust resistance gene Sr2 through marker‐assisted selection

Many stem rust resistance genes have been formally named in wheat. Adult plant stem rust resistance gene Sr2 was mapped in the short‐arm of chromosome 3B. Stripe rust resistance gene Yr57, identified in Aus91463, was mapped about 5 cM away from Sr2 based on its linkage with Sr2‐linked marker gwm533. The objective of this study was to combine Sr2 and Yr57 in a single genotype. A mapping population containing 107 recombinant inbred lines was developed from a cross between Aus91463‐Yr57 and Hartog‐Sr2. This population was tested at the seedling stage in the glasshouse for variation in stripe rust response, and high temperature induced Sr2‐linked seedling chlorosis. The RIL population was screened for Sr2‐linked pseudo black chaff phenotype at the adult plant stage in field. Five recombinants carrying Sr2 and Yr57 in coupling were detected using phenotypic and marker data. Four recombinants also carried leaf rust resistance gene Lr23 from Aus91463. These recombinants are being used as triple rust resistance source in the Australian Cereal Rust Control Program.     

Basic studies on hybrid wheat breeding using the 1BL-1RS translocation chromosome / Aegilops kotschyi cytoplasm system 1. Development of male sterile and maintainer lines with discovery of a new fertility-restorer

The Aegilops kotschyi cytoplasm and a 1BL-1RS translocation chromosome that consists of the long arm of wheat chromosome 1B and the short arm of rye chromosome 1R were transferred to six spring common wheat cultivars by repeated backcrossing. Resistance to leaf rust race 21B conditioned by the Lr26 gene and a secalin subunit encoded by the Sec-1 gene, both on the 1RS arm, were used as the selection markers of the translocation chromosome. Five of the six cultivars used were converted to complete male steriles, whereas the remaining one, cv. Kitamiharu 48, retained normal fertility, after transfer of both the 1BL-1RS chromosome and Ae. Kotschyi cytoplasm. Conventional gene analysis suggested that Kitamiharu 48 carries an incompletely dominant fertility-restoring gene. The F₁ hybrids between the male steriles and ordinary common wheat cultivars recovered fertility only at a low level, indicating that a single dose of the Rfv1 gene on the 1BS arm of wheat is insufficient for full fertility restoration under spring-sowing condition. Our results are in clear contrast to complete fertility restoration under fall-sowing condition reported by Nonaka et al. (1993). Combination of the 1BL-1RS chromosome / Ae. Kotschyi cytoplasm system with a new fertility-restoring gene discovered in Kitamiharu 48 may provide a breakthrough for spring-type hybrid wheat. This revised version was published online in August 2006 with corrections to the Cover Date.     

Rapid introduction of disease resistance from rye into common wheat by anther culture of a 6x triticale × nulli-tetrasomic wheat

Powdery mildew (caused by Erysiphe graminis) and yellow rust (caused by Puccinia striiformis) are the two most serious wheat diseases found in China. Rye chromosomes, carrying genes for resistance to these diseases, were introduced into common wheat in two generations using chromosome engineering and anther culture. The F₁ hybrids from a cross involving a hexaploid triticale (×Triticosecale Wittmack) בChinese Spring’ nulli‐tetrasomic N6DT6A wheat aneuploid line were anther cultured and doubled‐haploid plants were regenerated. Using genomic in situ hybridization, C‐banding and biochemical marker analyses, one of the anther‐cultured lines (ZH‐1)studied in detail, proved to be a doubled‐haploid with one rye chromosome pair added (1R) and a homozygous 6R/6D substitution (2n= 44). The line was tested for expression of disease resistance and found to be highly resistant to powdery mildew and moderately resistant to yellow rust.     

Identification of microsatellite markers for a leaf rust resistance gene introgressed into common wheat from Triticum timopheevii

The tetraploid wheat Triticum timopheevii Zhuk (A^tA^tGG) is known as a source of genes determining resistance to many diseases. An introgressive line 842, with durable resistance to leaf rust was established by crossing T. aestivum cv. ‘Saratovskaya29’ with T. timopheevii ssp. viticulosum and used for mapping leaf rust resistance genes. Molecular analysis of the line 842 with polymorphic microsatellite markers detected introgressions of T. timopheevii into the homoeologous group 2 chromosomes of common wheat. Transloca‐tion breakpoints of introgressed fragments were localized between the markers Xgwm95 and Xgwm817 on chromosome 2A, as well as Xgwm1128 and Xgwm1067 on chromosome 2B. Linkage analysis demonstrated the association of disease resistance at the seedling stage with chromosome 2A. The gene was found to be linked with marker Xgwm817 at a genetic distance of 1.5 cM. The alien leaf rust resistance gene was temporarily designated as lrTt1.     

Tan-Spot Resistance Screening of Aegilops Species

Two hundred and twelve accessions of 8 diploid and 10 polyploid species of Aegilops were evaluated for resistance to tan-spot disease of wheat, caused by Pyrenophora tritici-repentis (Died.) Drechs., under greenhouse conditions. One or more accessions of Ae. bicornis, Ae. biuncialis, Ae. Crassa, Ae. columnaris, Ae. cylindrica, Ae. speltoides, Ae. squarrosa. Ae. triaristata. Ae. triuncialis, and Ae. Ovata showed resistance following a 24-hour post-inoculation wet period. With a 36-hour wet period, diploids became only slightly or moderately susceptible and resistant polyploids became susceptible. A 48-hour wet period resulted in still greater susceptibility of both diploid and polyploid species.     

Utilisation of <Emphasis Type="Italic">Aegilops</Emphasis> (goatgrass) species to widen the genetic diversity of cultivated wheat

Wild Aegilops species related to cultivated wheat (Triticum spp.) possess numerous genes of agronomic interest and can be valuable sources of resistance to diseases, pests and extreme environmental factors. These genes can be incorporated into the wheat genome via intergeneric crossing, following, where necessary, the development of chromosome addition and substitution lines from the resulting hybrids. The transfer of a single segment from an alien chromosome can be achieved by translocations. The Aegilops (goatgrass) species, which are the most closely related to wheat, exhibit great genetic diversity, the exploitation of which has been the subject of experimentation for more than a century. The present paper gives a survey of the results achieved to date in the field of wheat–Aegilops hybridisation and gene transfer. The Aegilops genus consists of 11 diploid, 10 tetraploid and 2 hexaploid species. Of these 23 Aegilops species, most of the diploids (Ae. umbellulata Zhuk., Ae. mutica Boiss., Ae. bicornis (Forssk.) Jaub. & Spach, Ae. searsii Feldman & Kislev ex Hammer, Ae. caudata L., Ae. sharonensis Eig, Ae. speltoides Tausch, Ae. longissima Schweinf. & Muschl.) and several polyploids (Ae. ventricosa Tausch, Ae. peregrina (Hack. In J. Fraser) Marie & Weiller, Ae. geniculata Roth, Ae. kotschyi Boiss., Ae. biuncialis L.) have been used to develop wheat–Aegilops addition lines. Wheat–Aegilops substitution lines were developed using several species, including Ae. umbellulata, Ae. caudata, Ae. tauschii, Ae. speltoides, Ae. sharonensis, Ae. longissima and Ae. geniculata. Translocations carrying genes responsible for useful agronomic traits were developed with Ae. umbellulata, Ae. comosa, Ae. ventricosa, Ae. longissima, Ae. speltoides and Ae. geniculata. A large number of genes were transferred from Aegilops species to cultivated wheat, including those for resistance to leaf rust, stem rust, yellow rust and powdery mildew, and various pests (cereal cyst nematode, root knot nematode, Hessian fly, greenbug). Many molecular markers are linked to these resistance genes. The development of new molecular markers is also underway. There are still many untapped genetic resources in Aegilops species that could be used as resistance sources for plant breeding.     

Mapping of Lr56 translocation recombinants in wheat

In an attempt to transfer the Lr56/Yr38 resistance loci from Aegilops sharonensis to wheat, a 6A‐6S^sh chromosome translocation was produced. It involves essentially the entire chromosome 6S^sh with a small terminal segment of 6AL. Induced homoeologous recombination of the translocated chromosome with 6A produced numerous recombinants including three recombined chromosomes carrying Lr56 that could not be precisely mapped for lack of suitable markers. This study aimed to determine the chromosomal locations of the translocation breakpoints in these three recombinants using various DNA markers as well as physical and genetic mapping. The three recombinants Lr56‐39, ‐157 and ‐175 carry small segments of Ae. sharonensis chromatin distally to the Xgpw4329 and IWA5416 loci near the 6AS telomere. The Ae. sharonensis chromatin that remains in each line includes a homoeolocus of the wheat marker locus Xdupw217 (on 6BS) and its characteristic amplification product can be used as a dominant marker for the presence of Lr56. Of the three recombined chromosomes, Lr56‐157 retained the least alien chromatin and appears to be the best candidate for use in wheat breeding.     

Leaf rust and stripe rust resistance genes transferred to common wheat from Triticum dicoccoides

Linked leaf rust and stripe rust resistance genes introduced from Triticum dicoccoides protected common wheat seedlings against a range of pathotypes of the respective pathogens. The genes were chromosomally mapped using monosomic and telosomic analyses, C-banding and RFLPs. The data indicated that an introgressed region is located on wheat chromosome arm 6BS. The introgressed region did not pair with the ‘Chinese Spring’ 6BS arm during meiosis possibly as a result of reduced homology, but appeared to pair with 6BS of W84-17 (57% of pollen mother cells) and ‘Avocet S’. The introgressed region had a very strong preferential pollen transmission (0.96–0.98) whereas its transmission through egg cells (0.41–0.66) varied with the genetic background of the heterozygote. Homozygous resistant plants had a normal phenotype, were fertile and produced plump seeds. Symbols Lr53 and Yr35 are proposed to designate the respective genes.