Breeding oat for resistance to the crown rust pathogen Puccinia coronata f. sp. avenae: achievements and prospects.

Crown rust, caused by Puccinia coronata f. sp. avenae (Pca), is a significant impediment to global oat production. Some 98 alleles at 92 loci conferring resistance to Pca in Avena have been designated; however, allelic relationships and chromosomal locations of many of these are unknown. Long-term monitoring of Pca in Australia, North America and elsewhere has shown that it is highly variable even in the absence of sexual recombination, likely due to large pathogen populations that cycle between wild oat communities and oat crops. Efforts to develop cultivars with genetic resistance to Pca began in the 1950s. Based almost solely on all all-stage resistance, this has had temporary benefits but very limited success. The inability to eradicate wild oats, and their common occurrence in many oat growing regions, means that future strategies to control Pca must be based on the assumption of a large and variable prevailing pathogen population with high evolutionary potential, even if cultivars with durable resistance are deployed and grown widely. The presence of minor gene, additive APR to Pca in hexaploid oat germplasm opens the possibility of pyramiding several such genes to give high levels of resistance. The recent availability of reference genomes for diploid and hexaploid oat will undoubtedly accelerate efforts to discover, characterise and develop high throughput diagnostic markers to introgress and pyramid resistance to Pca in high yielding adapted oat germplasm.

First Report of a New TTKSF Race of Wheat Stem Rust (Puccinia graminis f. sp. tritici) in South Africa and Zimbabwe.

Seven races have been described in the Ug99 race group of Puccinia graminis f. sp. tritici (2). Ug99-related races previously recorded in South Africa are TTKSF, TTKSP, and PTKST (4). In December 2010, severe stem rust infection of the winter wheat cv. Matlabas was observed for the first time in South Africa. Race analysis using the 20 North American (NA) stem rust differential lines and letter code system classified the race as TTKSF. In comparative infection studies in a greenhouse, cv. Matlabas seedlings were susceptible (infection type [IT] 4) to isolate UVPgt61/1 (TTKSF+) collected from Afrikaskop in the eastern Free State, whereas the cultivar was resistant (IT 1 to 2) to stem rust isolates 2013 (TTKSF), UVPgt55 (TTKSF), UVPgt59 (TTKSP), and UVPgt60 (PTKST). Isolate 2013 represents the original collection of race TTKSF in South Africa (1). In addition to the NA differentials, no variation in the IT range of seedlings of lines with Sr7a, 8b, 12, 13, 14, 16, 18, 19, 22, 25, 26, 27, 28, 29, 32, 33, 34, 35, 39, 41, 42, 43, 44, Em, R, Tt2, and Satu was observed between UVPgt61/1 and UVPgt55. With the exception of cv. Matlabas, ITs of 106 South African cultivars likewise did not differentiate UVPgt61/1 and UVPgt55. Seedling IT studies were conducted at least twice. Microsatellite analysis (4) showed that all single pustule isolates established from the original Matlabas isolate formed part of the Ug99 group. When characterized with selected single nucleotide polymorphisms (SNPs), all single pustule isolates shared an identical genotype that differed from UVPgt55 (TTKSF), a foreign introduction into South Africa (1,3). SNP genotype analysis suggests that UVPgt61/1 is genetically dissimilar to UVPgt55, as is Zim1009, another TTKSF+ isolate that was collected from Birchenough in Zimbabwe. Studies are underway to determine the identity of the defeated Sr gene in Matlabas and the cultivar has been added to the South African stem rust differential set. TTKSF+ is the eighth race detected in the Ug99 group. Since no other cultivars or advanced lines were found to carry the Matlabas gene, it is unlikely that race TTKSF+ will threaten wheat production in South Africa. However, the occurrence of a new Ug99-related race emphasizes the variability within this internationally important group. References: (1) W. H. P. Boshoff et al. Plant Dis. 86:922, 2002. (2) R. F. Park et al. Euphytica 179:109, 2011. (3) B. Visser et al. Mol. Plant Pathol. 10:213, 2009. (4) B. Visser et al. Euphytica 179:119, 2011.

A genetic analysis of adult plant resistance to stripe rust in the wheat cultivar Kariega.

The wheat cultivar Kariega expresses complete adult plant resistance to stripe rust in South Africa. The aim of this investigation was to determine the extent and nature of variability in stripe rust resistance in a population of 150 doubled haploid lines generated from a cross between Kariega and the susceptible cultivar Avocet S. Analysis of field data for adult plant stripe rust resistance identified two major QTLs and two minor QTLs in the resistant cultivar Kariega. The two major QTLs were located on chromosomes 7D ( QYr.sgi-7D) and 2B ( QYr.sgi-2B.1), contributing 29% and 30% to the phenotypic variance, respectively. QYr.sgi-2B.1 is primarily associated with a chlorotic and/or necrotic response, unlike QYr.sgi-7D, which is believed to be the adult plant resistance gene Yr18. These two QTLs for adult plant resistance in Kariega appear to represent different forms of resistance, where QYr.sgi-7D may represent potentially more durable resistance than QYr.sgi-2B.1. Mixture model analysis of the field leaf infection scores suggested a genetic model involving two independent genes combining in a classical, epistatic manner. The results of the QTL analysis demonstrate its higher resolution power compared to the mixture model analysis by detecting the presence of minor QTLs.

First Report of Virulence in Puccinia graminis f. sp. tritici to Wheat Stem Rust Resistance Genes Sr8b and Sr38 in South Africa.

During the 2000 to 2001 season, 27 stem rust samples were collected from wheat (Triticum aestivum), barley (Hordeum vulgare), and triticale (× Triticosecale) cultivars and lines in the Western Cape, South Africa. Following inoculation and multiplication on McNair 701 seedlings, 40 single pustule isolates of P. graminis f. sp. tritici were established. Twenty-six isolates obtained from wheat, barley, or triticale that produced a similar reaction pattern on a set of differentiating host lines, were designated as pathotype Pgt-2SA55. Pgt-2SA55 is avirulent to Sr5, -6, -7b, -8b, -9b, -9e, -9g, -23, -24, -27, -30, -38, and -Gt, and virulent to Sr11, and -Agi. The remaining 14 isolates, all from wheat and designated as pathotype Pgt-2SA88, were avirulent to Sr24, -27, and -Agi, and virulent to Sr5, -6, -7b, -8b, -9b, -9e, -9g, -11, -23, -30, -38, and -Gt. On an expanded differential set, representative isolates of each pathotype were all avirulent to Sr13, -15, -21, -22, -25, -26, -29, -31, -32, -33, -35, -39, -43, and -Em, and virulent to Sr7a, -8a, -9a, -9d, -9f, -10, -12, -14, -16, -19, -20, -34, and Lc. Pgt-2SA55 was avirulent on cv. Renown (Sr2, -7b, -9d, and -17), whereas Pgt-2SA88 was virulent on this cultivar and Line R Sel carrying only Sr17. Both pathotypes differ from those identified previously in South Africa (1) and to our knowledge, Pgt-2SA88 is the first local isolate to have virulence towards Sr8b and the T. ventricosum-derived gene Sr38. Virulence to Sr38 has been reported in a P. graminis f. sp. tritici isolate collected in Uganda (2). Pathotype Pgt-2SA88 is virulent to seedlings of the previously resistant South African cvs. SST 57 (heterogeneous), Tugela, Tugela DN, and PAN 3377. Furthermore, 20% of the elite breeding lines in the spring and winter wheat breeding program of the Small Grain Institute expressed susceptible seedling reactions to Pgt-2SA88. Triticale cvs. Rex and Kiewiet were heterogeneous in their seedling reaction towards Pgt-2SA55. Seedling and field reactions recorded for the barley cvs. Sterling and SSG 532 and the experimental varieties Puma and Jaguar, showed an increase in stem rust susceptibility to Pgt-2SA55 when compared with existing South African pathotypes. The higher incidence of stem rust in commercial fields and experimental plots of wheat and barley in the Western Cape may be attributed to a recent increase in the cultivation of stem rust-susceptible cultivars in the region. The detection of two new pathotypes of P. graminis f. sp. tritici is of concern to the local small grain industry and requires continued resistance breeding. References: (1) W. H. P. Boshoff et al. S. Afr. J Plant Soil 17:60, 2000. (2) Z. A. Pretorius et al. Plant Dis. 84:203, 2000.

Establishment, Distribution, and Pathogenicity of Puccinia striiformis f. sp. tritici in South Africa.

Stripe rust, caused by Puccinia striiformis Westend. f. sp. tritici Eriks., has become an endemic disease of wheat (Triticum aestivum L.) in South Africa since it was first observed near Moorreesburg, Western Cape during August 1996. The main objectives of this study were to monitor the occurrence, spread, and the possible development of new variants of the stripe rust pathogen and the susceptibility of grass species to the pathogen. Results of surveys conducted during 1996 to 1999 revealed that rainfed wheat produced in the Western Cape, Eastern Cape, and the eastern Free State, as well as irrigated wheat produced in KwaZulu-Natal and the Free State, are most likely to be affected by stripe rust epidemics. Pathotype 6E16A- with virulence to Yr2, Yr6, Yr7, Yr8, Yr11, Yr14, Yr17, and Yr19 and pathotype 6E22A- with added virulence to Yr25 were detected. The occurrence of pathotype 6E22A- is currently restricted to KwaZulu-Natal and the Free State. Stripe rust isolates found on Hordeum murinum L. in the Western Cape were identified as pathotype 6E16A-, and both pathotypes 6E16A- and 6E22A- were collected from Bromus catharticus Vahl (= B. unioloides H.B.K.) in the eastern Free Sate. Urediospores from infections similar to stripe rust found on the grass species Dactylis glomerata L. (Eastern Cape), Poa pratensis L. (= P. bidentata Stapf; Western Cape), and P. annua and P. triviales L. (eastern Free State) failed to infect wheat cv. Morocco seedlings in the glasshouse. The possible role of grasses in the over-summering of the stripe rust pathogen has not yet been established. Stripe rust infections, however, have been found on summer-sown wheat in the south Western Cape during 1998, volunteer wheat growing in the summer and autumn months in the eastern Free State from 1998 to 2000, and on summer-sown wheat in Lesotho.

Occurrence and Pathogenicity of Puccinia hordei on Barley in South Africa.

Although leaf rust, caused by Puccinia hordei, is considered an important disease of barley (Hordeum vulgare) and regularly reaches epidemic proportions, pathogenic variability has never been studied in South Africa. From 1994 to 1997, only one pathotype (SAPh 3231) was identified with virulence to resistance genes Rph1, Rph4, Rph5, Rph10, and Rph11. During 1998, a second pathotype (SAPh 7321) was identified, differing from pathotype SAPh 3231 only in virulence to Rph12. Pathotype SAPh 7321 increased rapidly in the area where it was first detected, resulting in localized epidemic outbreaks in 1999. The reactions of various South African cultivars and breeding lines toward these pathotypes were determined, and the presence of Rph12 was postulated for B93/4, Krona, Optic, Prisma, and SSG 532. Rph genes showed varying degrees of temperature sensitivity, with none of the known genes displaying major changes in their phenotypes except Rph8, which was less effective at higher temperatures. Eight accessions of two wild Hordeum spp. occurring abundantly in the barley growing regions were found to be either weak or nonhosts for P. hordei.

Pathogenic Variability of Puccinia coronata f. sp. avenae and P. graminis f. sp. avenae on Oat in South Africa.

Although crown rust (caused by Puccinia coronata f. sp. avenae) and stem rust (caused by Puccinia graminis f. sp. avenae) are generally considered to be the most widespread and damaging diseases of oat (Avena spp.) in South Africa, pathogenic variability has never been studied. During 1997 and 1998, one dominant crown rust pathotype (SBLL) was identified with virulence to resistance genes Pc40, Pc45, Pc46, Pc51, and Pc54. Four other pathotypes (SGLL, PBBB+Pc35, SDQL, and JBBM+Pc35), occurring at low frequencies and further rendering resistance genes Pc35, Pc39, Pc48, Pc50, Pc52, and Pc64 ineffective, were also detected. Resistance gene Pc40 was postulated in Wisconsin X1588-2; Pc51 in Euro, Maluti, Overberg, OX88I 075-106, Perdeberg, and Swartberg; and Pc39 was confirmed in the cultivar Fidler. During the same period, four stem rust pathotypes were identified with virulence to resistance genes Pg1, Pg2, Pg4, Pg8, Pg9, Pg12,Pg15, and Pga. Resistance gene Pga was postulated in Alpha, OX87 080-1, OX88I 075-106, Sederberg, and W94/4; Pg2 and/or Pg4 in Euro, Perdeberg, Potberg, and Swartberg; and Pg9 in Pallinup and Victorian. Collections of wild oat species Avena fatua, A. byzantina, A. sterilis, and A. barbata were susceptible to all crown rust patho-types, while the four stem rust pathotypes were virulent on all species except A. barbata.

A New Pathotype of Puccinia striiformis f. sp. tritici on Wheat in South Africa.

Following the detection of Puccinia striiformis f. sp. tritici for the first time on wheat (Triticum aestivum) in the Western Cape in August 1996, stripe rust has spread to all the important wheat production areas in South Africa. Only the introduced pathotype (pt. 6E16) was detected in surveys of these areas during 1996 and 1997. In 1998, a severe stripe rust epidemic occurred in the eastern Free State on the extensively grown cultivars Hugenoot and Carina, both which are resistant to pt. 6E16. Stripe rust severities of 100% were common on flag and lower leaves, and widespread applications of fungicides were necessary. Avirulence/virulence characteristics of P. striifomis f. sp. tritici isolates collected from Hugenoot and Carina were determined on 17 standard stripe rust differential wheat lines and 11 supplementary testers. The latter testers included the wheat lines TP981 and TP1295 (supplied by R. Johnson, Cambridge, UK), both of which have a major resistance gene in common with the differentials Heines Peko, Reichersberg 42, Strubes Dickkopf, Clement, and Heines VII (1). Isolates obtained from Hugenoot and Carina differed from pt. 6E16 based on virulence to Reichersberg 42 (Yr7,25), Heines Peko (Yr2,6,25), TP981 (Yr25), and TP1295 (Yr25). The new variant, designated as 6E22, was also identified in collections from the province KwaZulu-Natal. Seedling tests with 6E16 and 6E22 have shown that Hugenoot, Carina, and Tugela-DN are the only local cultivars affected by the new pathotype. The occurrence of pt. 6E22, which appears to be a single-step adaptation from 6E16 adding virulence to Yr25, emphasizes the vulnerability of monogenic resistance to this disease. Reference: (1) R. A. McIntosh et al. Wheat Inform. Serv. 85:56, 1997.

First Report of Puccinia striiformis f. sp. tritici on Wheat in South Africa.

During August 1996, stripe (yellow) rust, caused by Puccinia striiformis f. sp. tritici, was observed for the first time on bread wheat (Triticum aestivum) in the Western Cape, South Africa. Ensuing surveys during the growing season indicated that stripe rust occurred throughout most of the wheat-producing areas in the winter rainfall regions of the Northern, Western, and Eastern Cape provinces. The disease was also observed on irrigated wheat in the summer rainfall area south of Kimberley. Stripe rust was most severe in the Western Cape, where prolonged cool and wet conditions favored epidemic development and necessitated extensive and often repeated applications of triazole fungicides. Due to spike infection and destruction of foliage, significant losses in grain quantity and quality occurred in certain fields. Avirulence/virulence characteristics of 32 stripe rust isolates, collected from commercial wheat fields, trap nurseries, and triticale, were determined on 17 standard differential wheat lines and seven supplementary testers supplied by C. R. Wellings, Plant Breeding Institute, Cobbitty, Australia. All isolates were representative of one pathotype, characterized by avirulence to Chinese 166 (Yr1), Vilmorin 23 (Yr3), Moro (Yr10), Strubes Dickkopf, Suwon 92/Omar, Clement (Yr2,9), Triticum aestivum subsp. spelta var. album (Yr5), Hybrid 46 (Yr4), Reichersberg 42 (Yr7), Heines Peko (Yr2,6), Nord Desprez (Yr3), Carstens V, Spaldings Prolific, Heines VII (Yr2), Federation*4/Kavkaz (Yr9), and Avocet-S/Yr15, and by virulence to Kalyansona (Yr2), Heines Kolben (Yr2,6), Lee (Yr7), Compair (Yr8), and Federation 1221. Cultivars Trident (Yr17), Avocet-R (YrA), and Selkirk (YrSk) appeared heterogeneous for stripe rust reaction. The pathotype resembled race 6E16, previously detected in East and North Africa, the Middle East, and western Asia. Pathotype identity was confirmed at IPO-DLO, Wageningen, using one South African isolate of P. striiformis f. sp. tritici. In view of the rapid dispersal of the pathogen during 1996, susceptibility of several high-yielding cultivars, and favorable climatic conditions in many wheat-growing areas, stripe rust is considered potentially damaging to South African wheat production. Field observations and seedling tests have shown, however, that certain cultivars are resistant to the introduced pathotype. At present the genetic basis of this resistance is largely unknown.