Transgressive segregation for resistance to yellow rust in wheat

Summary Crosses were made between wheat varieties Joss Cambier, Nord Desprez and Maris Bilbo, all classified as susceptible to yellow rust in field tests, and between Cappelle Desprez and Maris Huntsman, both classified as moderately and durably resistant. Selection for resistance to yellow rust among the progeny was carried out using races of Puccinia striiformis able to overcome all the known race-specific components of resistance in both parents of each cross. Lines with greater resistance than in both parents were obtained from each cross, those with greatest resistance being obtained from the cross between the moderately resistant parents. Three lines selected for resistance from the cross of Joss Cambier with Nord Desprez and one from the cross of Cappelle Desprez with Maris Huntsman, together with the parents, were tested in the field with 12 races of P. striiformis. Nord Desprez possessed a previously undetected race-specific component. The selected lines also displayed race-specific resistance, some of which was clearly related to race-specificity of the parents, and a component of resistance, greater than in both parents, that was effective against all 12 races. The possible origin and potential durability of this transgressive level of resistance is discussed. It is suggested that such transgressive resistance is more likely to be durable if it is derived from parents that have shown durable resistance.     

Genes for resistance to stripe rust in four spring wheat varieties

Summary Four spring wheat (Triticum aestivum L.) varieties differing in origin and reaction in the seedling stage to pathotype CDL-6 (extant in California) were intercrossed and examined in greenhouse conditions in F₁, F₂, and F₃ generations. Digenic and transgressive segregation was found in all crosses. The four varieties each had infection types (1 immune, 9 susceptible) and putative resistance genes as follows: Anza, IT 7, YrA; Glennson 81, IT 2, Yr9; Yecora Rojo, IT 6, YrC; and Ollanta, IT 4–6, YrL. Anza was classified as susceptible, Yecora Rojo and Ollanta as intermediate in seedling resistance, and Glennson 81 as resistant in the seedling stage.     

Postulation of resistance genes to yellow rust in wild emmer wheat derivatives and advanced wheat lines from Nepal

Summary Seedlings of 38 wild emmer derivatives, and a total of 53 advanced wheat varieties/lines introduced from the International Maize and Wheat Improvement Centre (CIMMYT) or other sources, Nepalese breeding lines and local cultivars were inoculated with 18 different yellow rust isolates to postulate yellow rust resistance genes (Yr). Many wild emmer wheat derivatives used were resistant to all isolates indicating the presence of undescribed genes. Some derivatives carried Yr9, Yr6 and/or YrSU. Genes Yr1, Yr2, Yr6, Yr7, Yr8, Yr15, YrSU and YrA+ are no longer effective in Nepal; Yr4, Yr5, Yr9, Yr10, YrSP and YrSD are still effective; the effectiveness of Yr3 remains unclear. This study shows that stripe rust resistance in seedling stage of most Nepalese cultivars and advanced materials is based on Yr9 with combinations of Yr2, Yr6, Yr7, and YrA+, of which only Yr9 is still effective in Nepal. In many countries Yr9 has lost its effectiveness. Therefore the introduction of new Yr-genes from wild emmer wheat in Nepalese cultivars is highly important.     

A recessive resistance gene for yellow rust (Puccinia striiformis West.) in bread wheat (Triticum aestivum L.)

Summary Three lines derived from the old dirty Dutch land variety Gelderse Ris were resistant against race 66(70)EO(16) of yellow rust. It was found that this resistance was conditioned by one recessive gene provisionally coded yrGR.     

Does interplot interference affect the screening of wheat for yellow rust resistance?

Summary In small plots, adjacent to one another, a representational error can be expected when screening for quantitative forms of resistance to airborne pathogens. The representational error or interplot interference may occur as an underestimation of the level of resistance and/or as an error in the ranking of the entries tested.Four experiments were carried out with wheat (Triticum aestivum) exposed to yellow rust (Puccinia striiformis), three in Kenya, one in Mexico (exp. III).In experiment I 57 entries were compared in an unreplicated trial with three adjacent and one isolated plot situation. The range in the standard deviation of and the ranking order for disease severity (DS) between the 57 entries were the same for all plot situations at all observation dates.In experiment II nine entries from experiment I, representing a wide range of quantitative resistance and having a similar heading date, were compared in adjacent plots consisting of two rows of 10 m in eight replicates and isolated plots of one, six and ten rows of 4 m in three replicates. The range in the standard deviation of and the ranking order for DS between the entries were very similar for the four plot situations.The 10 entries in experiment III differed from those in experiment II, but represented a similar wide range of resistance. Three adjacent plot situations of 0.9×0.5 m, 0.9×2.0 m and 2.7×2.0 m, respectively, were compared with one isolated one with plots of 2.7×2.0 m. The ranking order was not affected, the range in and the standard deviation of the DS in the isolated plots were slightly larger than in the adjacent plots.In experiment IV two mixtures of two entries each were made. Per mixture one entry was fairly resistant (R) the other rather susceptible (S). Within each mixture the entries had a similar heading date. The ratios of the R:S mixtures were 0:100, 50:50, 67:33, 75:25, 82:18 and 100:0. The DS of each entry was the same as its DS in monoculture irrespective of the mixture ratio. There was no mixture effect on DS.The three experiments in Kenya gave no indication of any interplot interference occurring. In Mexico there was a very slight underestimation of the resistance in adjacent plots. The ranking order was always the same irrespective of the test plot situation. The screening of wheat for yellow rust resistance in small adjacent plots is representative for the farmers fields. This is contrary to what has been found in other windborne pathosystems such as barley-barley leaf rust (strong under estimation of resistance), barley-powdery mildew (some under estimation of resistance and different ranking order) and durum wheat-stem rust (fair under estimation of resistance).     

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.     

Genes for resistance to stripe rust in four spring wheat varieties

Summary The stripe rust resistant spring wheat (Triticum aestivum L.) varieties Anza, Glennson 81, Ollanta, and Yecora Rojo gave 1,2,2, and 2-gene segregations, respectively, in hybrids with susceptible Jupateco 73 when inoculated in field conditions at Davis, California USA with Puccinia striiformis West. pathotype CDL-6 and rated at post-heading stage. Intercrosses of these varieties, Anza/Yecora Rojo was not studied, permitted the following conclusions about the genes expressed in adult plants: Anza, one recessive gene; Glennson 81, two dominant genes; Ollanta, two genes, at least one is dominant; and Yecora Rojo; one dominant and one recessive gene, one of which is common with Ollanta. The resistance genes in these varieties, which expressed resistance in the seedling stage, were believed to be effective at the adult stage. Thus, seven resistance genes were identified in the four varieties. The genotypes were designated for the purposes of this study as follows: Anza, YrA YrH; Glennson 81, Yr9, YrJ, Ollanta YrL YrD; and Yecora Rojo, YrC YrD. It was recommended that these and other Yr genes be used as multiple gene complexes to increase durability of resistance to P. striiformis, an organism known to evolve virulence rapidly in field conditions. The demonstrated durability of Anza in California may be a result of its combination of resistance alleles at two loci.     

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.     

Characterization of the durable resistance to yellow rust in old winter wheat cultivars in the Netherlands

Summary Winter wheat cultivars released in the Netherlands before 1930 carried durable resistance to yellow rust. Cultivars released in the period between 1930 and 1950 often were durably resistant while recent cultivars infrequently showed durable resistance. This durable resistance was not difficult to transfer to new cultivars. Twenty nine older cultivars with durable resistance and eight recent non-durably resistant cultivars were tested in the seedling stage and in the adult plant stage against 12 West-European yellow rust races and against some non-European races in the seedling stage only. The adult plant tests were carried out in race nursery tests in the Flevopolder. Per race nursery all 37 cultivars, planted in hills of about 20 plants on both sides of the highly susceptible cv. Michigan Amber, were exposed to one race.The infection type of each cultivar-race combination was scored on 0 to 9 scale once in the seedling stage and twice in the adult plant stage. In the race nurseries the percentage leaf area affected was evaluated three times to be used to calculate the area under the disease progress curve (AUDPC). This AUDPC multiplied with the mean infection type in the field gave the susceptibility index (SI).The infection types were classified into resistant (R), intermediate (I) or susceptible (S) when the infection types were 0 to 3, 4 to 6 or 7 to 9, respectively. Four categories of resistance were discerned on the basis of the three infection type scores: 1) RRR, overall resistance; complete or near-complete resistant at all stages. 2) SRR, adult plant resistance, complete- or near-complete resistant at the adult plant stage only. 3) SRS and SSR, temperature sensitive resistance, the resistance changed from the one evaluation data to the other. 4) SSS and an SI lower than that of Michigan Amber, partial resistance.The frequencies of overall, adult plant and temperature sensitive resistance were 1.4, 52 and 54% in the older cultivars and 40, 62 and 22% in the recent ones, respectively. Among the older cultivars all had a fair to high level of partial resistance, the SI being on average only 20% of that of Michigan Amber, while most cultivars also seemed to carry temperature sensitive resistance. The partial resistance of the recent cultivars was of a much lower level with a mean SI compared to that of Michigan Amber of 61%. Partial resistance was highly correlated (r = –0.94) with the mean resistance scores from the Dutch Recommended Cultivars Lists. It was concluded that partial resistance and temperature sensitive resistance were the major components of the durable resistance in the older cultivars.     

Evolution of virulence patterns in yellow rust races and its implications for Ereeding for resistance in wheat in Kenya

Summary Virulence patterns of yellow rust isolates collected in Kenya between 1986–1989 were compared with earlier results. The number of virulence factors per race and the range in virulence factors both increased considerably. Before 1976 races carried on average 4.5 to 5.0 virulence factors, whereas the races after 1986 had a mean of 6.5 virulence factors. The range in the number of virulence factors increased from some seven to eight in the first period to 12 in the second out of the 17 evaluated. In the period 1986–1989 another three virulence factors (2, 9 and A) were assessed. All three occurred at a high frequency.Virulence neutralizing the resistance genes Yr2, Yr2+, Yr6, Yr6+, Yr7, Yr7+, Yr8, Yr9, Yr9+ and those in the cultivars Anza (A), Strubes Dickkopf (SD) and Suwon92/Omar (SU) occurred at a high frequency, while virulence for Yr3V, Yr4+, Yr5, CV and SP (resistance in Carstens V and Spaldings Prolific resp.) were not found. The remaining three virulence factors for Yr1, 10 and 3N were rare.In the past ten years the resistance of most released cultivars became ineffective in less than six years. They were shown to carry race-specific major resistance genes such as Yr7+, Yr9+, SD and A. However, in the field, the resistance of the cultivars was not completely neutralized. A residual resistance, ranging from moderate to fairly high, was observed in all cultivars in which the major gene resistances were neutralized by corresponding virulence genes.Other wheat cultivars such as Africa Mayo, Kenya Kudu, Enkoy, Kenya Leopard, Bounty, Frontatch, Bonny and Kenya Plume appeared to keep their resistance over a condiserable period of time. They are considered to be durably resistant to the Kenyan yellow rust populations. This form of resistance, together with the residual resistance, can be recommended for use in breeding programmes.     

Resistance to stripe rust in Triticum turgidum,T. tauschii and their synthetic hexaploids

Resistance to stripe rust (caused by Puccinia striiformis Westend.) of 34 Triticum turgidum L. var.durum, 278 T. tauschii, and 267 synthetic hexaploid wheats (T. turgidum x T. tauschii) was evaluated at the seedling stage in the greenhouse and at the adult-plant stage at two field locations. Mexican pathotype 14E14 was used in all studies. Seedling resistance, expressed as low infection type, was present in all three species. One hundred and twenty-eight (46%) accessions of T. tauschii, 8 (23%) of T. turgidum and 31 (12%) of synthetic hexaploid wheats were highly resistant as seedlings. In the field tests, resistance was evaluated by estimating area under disease progress curve (AUDPC). Synthetic hexaploid wheats showed a wide range of variability for disease responses in both greenhouse and field tests, indicating the presence of a number of genes for resistance. In general, genotypes with seedling resistance were also found to be resistant as adult plants. Genotypes, which were susceptible or intermediate as seedlings but resistant as adult plants, were present in both T. turgidum and the synthetic hexaploids. Resistances from either T. turgidum or T. tauschii or both were identified in the synthetic hexaploids in this study. These new sources of resistance could be incorporated into cultivated hexaploid wheats to increase the existing gene pool of resistance to stripe rust.     

Stripe rust reactions of Australasian wheats

Summary Seedling and field reactions to four European races of stripe rust were determined for 254 wheat cultivars, mostly from Australia and New Zealand. Numerical analyses of the data employed a divisive classification procedure with termal reallocation to form 10 groups which were then further classified, ordinated and diagnosed by several procedures. The groups formed ranged from one comprising four wheats with low seedling reactions to all four rust strains and mean field rust of only 1.0%, to one comprising 105 wheats with high seedling reactions and mean field rust of 64.7%. Groups of cultivars with intermediate levels of rust resistance were classified both on relative level of overall rust and on differential reactions to rust strains. One group of 13 cultivars had moderately high seedling reactions but averaged only 1.5% rust in the field.Seedling response to gibberellic acid (GA) was also measured and diagnosed as an external attribute, along with grain type and region of origin. Principal co-ordinate analysis revealed that greater resistance to stripe rust was associated with the insensitivity to GA typical of semidwarf wheats and with an origin in northern Australia, where semidwarf wheats have been most used in the breeding program. In terms of numerical analysis of disease data in plant breeding programs, the study highlighted the advantages of using divisive classification with terminal reallocation of group members.     

Inheritance of slow rusting to stem rust in wheat

Summary The inheritance of the slow rusting character was studied on F₅ progenies from seven spring wheat cultivars (Triticum aestivum) crossed in all possible combinations without reciprocals. The cultivars and their progenies were evaluated for slow rusting in 1974 and 1975 in epidemics of Puccinia graminis f. sp. tritici, races 15 and 151, and traces of other races. Slow rusting varied significantly among the parents and among the F₅ progeny of each cross. Transgressive segregation occurred in each cross, i.e. some progeny rusted more slowly than the parents and some faster. In crosses with both Idaed 59 and Kenya 58 the progeny distributions were skewed towards slow rust development but the distributions in the other crosses were normal. The genetic control of slow rusting was predominantly additive, and narrow sense heritability was approximately 80 percent. The number of segregating genes having an effect on slow rusting was estimated to be 2 to 12 pairs depending on the cross. Correlation between slow rusting and maturity was usually negative but in most crosses the relationship was small.Contribution No. 9624 from the Agricultural Experiment Station, University of Minnesota, St. Paul, Minnesota 55108.     

Cytogenetical studies in wheat XIX. Location and linkage studies on gene Yr27 for resistance to stripe (yellow) rust

The stripe (yellow) rust resistance gene Yr27 was located in wheat (Triticum aestivum L.) chromosome 2B and shown to be closely linked to the leaf (brown) rust resistance genes Lr13 and Lr23 in the proximal region of the short arm. Gene Yr27 was genetically independent of Lr16, which is distally located in the same arm. While Yr27 was often difficult to score in segregating seedling populations, it is apparently quite effective in conferring resistance to avirulent cultures under field conditions. The occurrence of Yr27 in Mexican wheat germplasm and the current over-dependence on Yr27 for crop protection in Asia are discussed.     

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

Summary The 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.     

Evaluation of seedling and adult plant resistance to leaf rust in European wheat cultivars

Summary A set of 105 European wheat cultivars, comprising 68 cultivars with known seedling resistance genes and 37 cultivars that had not been tested previously, was tested for resistance to selected Australian pathotypes of P. triticina in seedling greenhouse tests and adult plant field tests. Only 4% of the cultivars were susceptible at all growth stages. Twelve cultivars lacked detectable seedling resistance to leaf rust, and among the remaining cultivars, 10 designated genes were present either singly or in combination. Lr13 was the most frequently detected gene, present in 67 cultivars, followed by the rye-derived gene Lr26, present in 19 cultivars. Other genes present were Lr1, Lr3a, Lr3ka, Lr10, Lr14a, Lr17b, Lr20 and Lr37. There was evidence for unidentified seedling resistance in addition to known resistance genes in 11 cultivars. Field tests with known pathotypes of P. triticina demonstrated that 57% of the cultivars carried adult plant resistance (APR) to P. triticina. The genetic identity of the APR is largely unknown. Genetic studies on selected cultivars with unidentified seedling resistances as well as all of those identified to carry APR are required to determine the number and inheritance of the genes involved, to determine their relationships with previously designated rust resistance genes, and to assess their potential value in breeding for resistance to leaf rust.     

Genetic and environmental variability in wheat characteristics reported to be involved in morphological resistance to wheat stem rust

Summary Four morphological characteristics of wheat peduncles were evaluated for genetic and environmental variability. The thickness of the epidermis and the amount and distribution of chlorenchyma tissue in the peduncle were measured. The genetic variability was estimated to be large, whereas the environmental variability was small. It was also concluded that cultivars could be screened for morphological characteristics of the peduncle that contributed to non-specific resistance to stem rust with a minimum sample size of about 10 plants per replication from a single location.Paper No 9328, Scientific Journal Series, Minnesota Agricultural Experiment Station. University of Minnesota. St. Paul. MN 55108.     

The inheritance of stem rust and leaf rust resistance in some Ethiopian wheat collections

Summary Common and durum wheat populations obtained from Sweden and originally collected in Ethiopia were screened for resistance to steum rust and leaf rust. Resistant selections of common wheat were crossed and backcrossed with either stem rust susceptible RL6071, or leaf rust susceptible Thatcher. Genetic studies, based largely on tests of backcross F₂ families, showed that four of the selections had in common a recessive gene SrA. Plants with this gene were resistant (1⁺ infection type) to all stem rust races tested. This gene was neither Sr26 nor Sr29. The resistance of other selections, based on tests with an array of rust isolates, was due to various combinations of Sr6, 8a, 9a, 9d, 9c, 11, 13, 30, and 36. One of the selections had linked genes, Lr19/Sr25. Another selection had a dominant gene for resistance (;1 infection type) to all the races of leaf rust. With the possible exception of this gene for leaf rust resistance and SrA, no obviously new resistance was found.     

Variation of characters in near-isogenic lines of wheat with added genes for leaf rust resistance

Summary Using the cultivar Arina as the recurrent parent, six backcrosses were made with two donor lines carrying the leaf rust resistance genes Lr1 and Lr9, respectively. Selection for leaf rust resistance occurred at the seedling stage in the greenhouse; the first plants transferred to the field were BC₆F₄s. Frequency distribution of the 332 Lr1/7 × Arina and the 335 Lr9/7 × Arina lines showed continuous variation for yellow rust resistance and heading date in these leaf rust near-isogenic lines (NILs). Similar results were also obtained for plant height, for resistance to powdery mildew and glume blotch, as well as for baking quality characters in another set of more advanced NILs. The available information on the behaviour of one of the parents of cultivar Arina led to the conclusion that the expressed yellow rust resistance is quantitative and might possibly be durable.     

Environmental stability of partial resistance in spring wheat to wheat leaf rust

Summary Five spring wheat cultivars differing in partial resistance (PR) to wheat leaf rust were tested at Wageningen (the Netherlands) on a sandy and a clay site, El Batan (CIMMYT, Mexico) and Ponta Grossa (Brazil) over two years. The cultivars were Skalavatis 56, Little Club (both very susceptible), Westphal 12A, Akabozu and BH 1146 (all three with high levels of PR). The results showed that PR was expressed at all four locations in both years. The level of expression was influenced by the environment but the cultivar ranking was hardly affected. Selection for PR in the field can therefore be carried out over a wide range of environments.