R. A. McIntosh

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

SummaryWild 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.

The Gene Sr33, an Ortholog of Barley Mla Genes, Encodes Resistance to Wheat Stem Rust Race Ug99

Resistance May Not Be Futile Recently, Ug99, a particularly devastating strain of wheat stem rust fungus, has emerged, which could potentially threaten food security. Now, two genes have been cloned that offer resistance to Ug99. Saintenac et al. (p. 783, published online 27 June) cloned Sr35 from Triticum monococcum, a diploid wheat species not often cultivated. Periyannan et al. (p. 786, published online 27 June) cloned Sr33 from Aegilops tauschii, a diploid wild grass that contributed to the hexaploid genome of cultivated wheat. The genes both encode proteins that show features typical of other disease resistance proteins and offer opportunities to slow the pace of Ug99 progression. Two resistance genes are identified that could protect wheat from a virulent fungus that can severely reduce crop yields. Wheat stem rust, caused by the fungus Puccinia graminis f. sp. tritici, afflicts bread wheat (Triticum aestivum). New virulent races collectively referred to as “Ug99” have emerged, which threaten global wheat production. The wheat gene Sr33, introgressed from the wild relative Aegilops tauschii into bread wheat, confers resistance to diverse stem rust races, including the Ug99 race group. We cloned Sr33, which encodes a coiled-coil, nucleotide-binding, leucine-rich repeat protein. Sr33 is orthologous to the barley (Hordeum vulgare) Mla mildew resistance genes that confer resistance to Blumeria graminis f. sp. hordei. The wheat Sr33 gene functions independently of RAR1, SGT1, and HSP90 chaperones. Haplotype analysis from diverse collections of Ae. tauschii placed the origin of Sr33 resistance near the southern coast of the Caspian Sea.

Mapping of durable adult plant and seedling resistances to stripe rust and stem rust diseases in wheat

Doubled haploid populations of CD87/Katepwa, Cranbrook/Halberd, and Sunco/Tasman were assessed for seedling response to stem rust and stripe rust. The CD87/Katepwa population was also screened as adult plants in the field against stripe rust. The respective parents differed in presence or absence of various stem rust and stripe rust resistance genes. At least 4 resistance loci controlled adult plant resistance to stripe rust in the CD87/Katepwa population, and based on quantitative trait loci mapping results, two of these were contributed by CD87. Pedigree information indicated that these regions correspond to durable adult plant stripe rust resistance genes Yr18 and Yr29. Yr29 was mapped to the distal region of chromosome 1BL. The third gene, contributed by Katepwa, YrKat, was located in chromosome arm 2DS. Sr30 mapped distal to markers abg3 and P36/M61-170 in chromosome arm 5DL. Genes Yr7 and Pbc (completely linked with durable stem rust resistance gene Sr2) showed close associations with markers in chromosome arms 2BL and 3BS, respectively. A distally located genomic region in chromosome 6AS also affected the expression of Pbc. The temperature-sensitive stripe rust resistance gene, YrCK, carried by Sunco showed monogenic inheritance and was located in chromosome arm 2DS. Several markers showed complete association with Triticum timopheevi derived stem rust resistance gene Sr36. Microsatellite markers stm773 and gwm271A were validated on a set of wheat genotypes and were found to be diagnostic for the detection of Sr36. TheSr36-linked Xstm773 allele showed better amplification than the Sr36-linked Xgwm271A allele. These markers could be used for marker assisted identification of Sr36 in breeding populations.

Cytogenetical studies in wheat

SummaryTwo genes for resistance to Puccinia graminis f. sp. tritici were located in chromosome 2B of Kota wheat. Both were mapped in the long arm. The first designated Sr28 was situated distally to Sr9 and showed 34·6±2·8 per cent recombination with the centromere. The second gene, which was not definitely distinguished from Sr16 either on the basis of recombination or by its response to several pathogen cultures, was inherited independently of the 2B centromere and of Sr9, but showed 38·2±1·9 and 29·2±4·2 per cent recombination with Sr28. Comparisons of present results with those from earlier studies suggested that Kota possesses at least five genes for resistance.

Linkage mapping of genes for resistance to leaf, stem and stripe rusts and ω-secalins on the short arm of rye chromosome 1R

SummaryThe genes controlling resistance to three wheat rusts, viz., leaf rust (Lr26), stem rust (Sr31) and stripe or yellow rust (Yr9), and ω-secalins (Sec1), located on the short arm of rye chromosome 1R, were mapped with respect to each other and the centromere. Analysis of 214 seeds (or families derived from them) from testcrosses between a 1BL.1RS/1R heterozygote and ‘Chinese Spring’ ditelocentric 1BL showed no recombination between the genes for resistance to the three rusts, suggesting very tight linkage or perhaps a single complex locus conferring resistance to the three rusts. The rust resistance genes were located 5.4 ± 1.7 cM from the Sec1 locus, which in turn was located 26.1 ± 4.3 cM from the centromere; the gene order being centromere — Sec1 — Lr26/Sr31/Yr9 — telomere. In a second test-cross, using a different 1BL.1RS translocation which had only stem rust resistance (SrR), the above gene order was confirmed despite a very large proportion of aneuploids (45.8%) among the progeny. Furthermore, a map distance of 16.0 ± 4.8 cM was estimated for SrR and the telomeric heterochromatin (C-band) on 1RS. These results suggest that a very small segment of 1RS chromatin is required to maintain resistance to all three wheat rusts. It should be possible but difficult to separate the rust resistance genes from the secalin gene(s), which are thought to contribute to dough stickiness of wheat-rye translocation lines carrying 1RS.

Validation of molecular markers for wheat breeding

Five sets of markers were assessed for their usefulness in breeding, two linked to wheat stem rust gene Sr2, several markers linked to a chromosome segment conferring Yr17/Lr37/Sr38 resistance, two reported markers for the linked genes Lr35 andSr39, one for Lr28, and one linked to flour colour. The gene for Sr2 confers adult plant resistance to stem rust (Puccinia graminis f.sp. tritici) and was originally transferred to bread wheat from the tetraploid emmer (‘Yaroslav’) to the cultivars Hope and H-44. The gene is located on the short arm of chromosome 3B and confers a durable adult plant resistance to stem rust usually expressed only in the field. The chromosome segment carrying the Lr37, Sr38, Yr17 resistance genes is located on 2AS and was originally introduced into wheat through an Aegilops ventricosa Triticum persicum cross, followed by a cross to the cultivar Marne (VPM1). The flour colour quantitative trait locus was originally described in a Yarralinka Schomburg cross and is located on chromosome 7A. The primers as originally developed required optimisation for more routine use in a breeding program.

Transfer of rye chromosome segments to wheat by a gametocidal system

A gametocidal chromosome derived from Aegilops triuncialis (3C) induces chromosome mutations in gametes lacking the 3C chromosome in common wheat (Triticum aestivum L.). We combined 3C with chromosome 1R of rye (Secale cereale L.) in a common wheat line to know how efficiently 3C induces transfers of small 1R segments to wheat. In the 811 progeny of this wheat line, we found five wheat chromosomes (2A, 2D, 3D, 5D and 7D) carrying segments of the 1R satellite. Wheat plants carrying these translocations were tested for the presence of a storage protein locus Sec-1 and a cluster of resistance genes for wheat rust diseases, Sr31, Lr26 and Yr9. The 2A and 2D translocations had the Sec-1 and three rust resistance loci. The 3D and 5D translocations had Sr31, Lr26 and Yr9 but not Sec-1. The 7D translocation lacked Sec-1, Lr26 and Yr9, but the presence of Sr31 in this translocation was not determined. This showed that the translocation points fell into three regions of the 1R satellite, namely, proximal to Sec-1, between Sec-1 and the rust resistance loci, and distal to the rust resistance loci. Thus, the 3C gametocidal system was demonstrated to be effective in transferring small rye chromosome segments.

Characterization of rust-resistant wheat-Agropyron intermedium derivatives by C-banding, in situ hybridization and isozyme analysis

SummaryChromosome constitutions of three wheat-Agropyron intermedium derivatives were identified by C-banding analysis, in situ hybridization using biotin-labeled genomic Ag. intermedium DNA as a probe and isozyme analysis. Lines W44 and W52 were identified as 7Ai-2(7D) and 7Ai-2(7A) chromosome substitution lines carrying the same chromosome pair of Ag. intermedium. The alien chromosome was found to be homoeologous to group 7 based on C-banding, meiotic pairing and isozyme analyses. Line W49 was identified as a wheat Ag. intermedium chromosome translocation line. The breakpoint of the T2AS · 2AL-7Ai-2L translocation is located in the long arm at a fraction length of 0.62, and the transferred Ag. intermedium segment has a size of about 2.4 μm. Lines W44 and W52 expressed Ag. intermedium genes for resistance to leaf rust, stripe rust and stem rust, but only leaf rust resistance was expressed in W49. The results show that the leaf rust resistance gene(s), designated Lr38, is located in the distal half of the long arm of chromosome 7Ai-2, whereas the genes for resistance to stem rust and stripe rust are located either in the short arm or in the proximal region of the long arm of this chromosome.

Postulation of leaf (brown) rust resistance genes in 70 wheat cultivars grown in the United Kingdom

Multi-pathotype tests on 70 U.K. wheat cultivars permitted postulation of eight known seedling genes for resistance to Puccinia recondita f. sp.tritici either singly or in combinations. The most commonly detected gene was Lr13 (present in approximately 57% of cultivars), followed by Lr26 (22%), Lr37 (20%), Lr10 (17%), Lr17b (LrH) (10%), Lr1 (7%), Lr3a (6%) and Lr20(4%). This information permitted assessments of adult plant resistance (APR) in some cultivars, in field nurseries inoculated with pathotypes of P. recondita f. sp. tritici of known pathogenicities for characterized seedling resistance genes. APR was identified in eleven cultivars, including Avalon and Maris Ranger, which lacked detectable seedling resistance genes. The results provided a better understanding of specific resistances in the cultivars tested than was available from previous reports.

Characterisation and origin of rust and powdery mildew resistance genes in VPM1 wheat

SummaryThe expression of rust resistances conferred by closely linked genes derived from VPM1 varied with environmental conditions and with genetic backgrounds. Under low light and low temperature conditions seedlings carrying Yr17 showed susceptible responses. Stem rust and leaf rust resistance genes Sr38 and Lr37 tended to confer more resistance at 17±2° C than at normal temperatures above > 20° C. These studies supported the hypothesis that Yr17, Lr37 and Sr38 were derived from Aegilops ventricosa, whereas Pm4b was probably derived from T. persicum. Studies on certain addition lines and parental stocks indicated that wheat cytoplasm may enhance the expression of Sr38.