Inheritance of the Powdery Mildew Resistance Gene Pm9 in Relation to Pm1 and Pm2 of Wheat

The inheritance of the powdery mildew resistance gene Pm9 originating from the hexaploid spring wheat cultivar ‘Normandie’ was analyzed in relation to Pm1 and Pm2. Two leaf segments of individual P_1?, P_2?, F_1? and F₂-plants of the cross ‘Normandie’ (Pm1, 2, 9) בFederation’ (no known Pm gene) were inoculated separately with two powdery mildew isolates. Using powdery mildew isolate No. 6 virulent for Pm1 and Pm2 but avirulent for Pm9, a 1 resistant (r): 3 susceptible (s) F₂-segregation was found for the Pm9 gene. Using powdery mildew isolate No. 3 virulent for Pm1 and Pm9 but avirulent for Pm2, a 3 (r): 1 (s) F₂-segregation was found for the Pm2 gene. Combining the data of both experiments (leaf segments of identical plants had been used), a 9 (sr): 3 (ss): 3 (rr): 1 (rs) segregation resulted for the F₂ of this cross: therefore, independent inheritance of the genes Pm2 and Pm9 can be concluded. Similarly, the cross ‘Mephisto’ (Pm1, 2, 9) בAmor’ (no known Pm gene) was analyzed. The Pm9 gene again showed a monogenically recessive inheritance, whereas Pm1 showed a monogenically intermediate segregation upon inoculation with powdery mildew isolate No. 9a virulent for Pm2 and Pm9 but avirulent for Pm1. Combining the single gene segregations, linkage between both genes was found among the progenies. A distance of 8.5 cM was calculated. Analyzing a set of spring wheat cultivars with seven defined powdery mildew isolates, the presence of Pm1, Pm2 and Pm9 in these lines was verified; in most cases, Pm1 occurred together with Pm9.     

New data on the specificity of the root-knot nematode resistance genes Me1 and Me3 in pepper

Root‐knot nematodes, Meloidogyne spp., cause severe damage on solanaceous crops, including tomato and pepper. The objective of this study was to test whether nematodes virulent against the tomato Mi resistance gene were able to overcome the Me1 and Me3 resistance genes from pepper. For that purpose, a collection of 22 Meloidogyne arenaria, Meloidogyne incognita and Meloidogyne javanica isolates, avirulent or virulent against the Mi gene, was assembled. The reproduction of each isolate was evaluated on both susceptible and resistant tomato and pepper genotypes in a growth chamber. The Me1 resistance gene controlled all the avirulent and Mi‐virulent nematodes tested, and therefore appears promising for pepper breeding. In contrast, one M. arenaria and two M. incognita virulent isolates were shown to overcome the pepper Me3 resistance gene. These results encourage the testing of a large number of Meloidogyne isolates to evaluate the plant‐nematode interaction, which could have important consequences for the use and management of resistance genes in the field with respect to specificity and durability of the resistance.     

Two unnecessary powdery mildew resistance genes in a synthetic rye population are neutral on fitness

A synthetic winter rye population was produced with two race-specific powdery mildew resistance genes, one dominant (Rm1) and the other (rm2) recessive, each at a frequency of about 0.50. The population was advanced by open-pollination in an isolated plot under mildew-free conditions for eight years. Samples of generations Syn-0 through Syn-7 were inoculated in the laboratory with two mildew isolates, one avirulent to either resistance gene, the other virulent to Rm1 and avirulent to rm2, to discriminate resistant and susceptible phenotypes. From the proportions of resistant plants, frequencies of Rm1 and rm2 were calculated and the fitness of carriers of resistance alleles was estimated in relation to carriers of susceptibility alleles at the two loci using continuous models and linear regression analyses. Frequencies of the two resistance genes oscillated only weakly over the eight generations. Coefficients of selection against Rm1-and rm2rm2 genotypes were –0.04 and –0.02, respectively, and not significantly different from zero. Thus the two resistance genes were selectively neutral. It is concluded that pyramiding of major powdery mildew resistance genes in rye varieties should not reduce their yield potential in the absence of mildew.     

Genes for race-specific resistance against blackleg disease in Brassica napus L.

Specificity of interaction at the cotyledon stage was recently demonstrated between the blackleg pathogen, Leptosphaeria maculans, and Brassica napus. Three pathogenicity groups were distinguished, PG2 avirulent towards ‘Quinta’ and ‘Glacier’, PG3 avirulent towards ‘Quinta’, and PG4 virulent on the two cultivars. The genetic control of the interactions was investigated on both the pathogen and the plant. Tetrad analysis was performed following PG3 × PG4 and PG2 × PG4 crosses.‘Quinta’ and ‘Glacier’ were crossed with the susceptible winter oilseed rape cultivar ‘Score’. The analysis of F₁, F₂ and testcross populations suggested that the incompatible interaction between ‘Quinta’ and PG3 isolates is conditioned by the presence of the dominant single resistance allele Rlml in ‘Quinta’ and the matching avirulence gene AvrLml in L. maculans. Race-specific resistance of ‘Glacier’ to PG2 isolates was conditioned by the matching gene pair Rlm2/AvrLm2. Finally, the data suggest that two avirulence genes matching two dominant loci control the ‘Quinta’-PG2 interaction. The consequences of the occurrence of race-specific resistance in B. napus are discussed with respect to future breeding for blackleg resistance.     

Inheritance of resistance to the bipartite Tomato chlorotic mottle begomovirus derived from Lycopersicon esculentum cv. ‘Tyking’

Severe outbreaks of bipartite begomoviruses (family Geminiviridae) have been observed on tomatoes after the introduction of the whitefly Bemisia tabaci (biotype B) in Brazil. The Lycopersicon esculentum line ‘TX 468-RG’ was identified as one of the best sources of broad-spectrum resistance to species comprising the tomato-infecting Begomovirus complex in Brazil. The genetic basis of resistance to one Begomovirus isolate was investigated using populations from the cross between ‘TX 468-RG’ (P₁) and the susceptible line ‘Ohio 8245’ (P₂). Parental lines, F₁, backcross (BC) to P₁ and BC to P₂ and F₂ generations were inoculated at the two true-leaf stage using 20 viruliferous whiteflies per plant. Assessment was done two weeks after inoculation based upon visual analysis of symptom expression. The ratio of resistant to susceptible plants closely fit to a single recessive gene (locus) model. The sequence analysis indicated that the Begomovirus isolate used in this assay was closely related to the bipartite Tomato chlorotic mottle virus. Therefore, this gene/locus, was tentatively named tcm-1 (tomato chlorotic mottle virus resistance-1). This locus has been transferred to distinct tomato cultivars and levels of resistance similar to that of ‘TX 468-RG’ were observed in advanced (F₈ and F₉) generations. In addition, breeding lines carrying the tcm-1 locus were also resistant to other Brazilian bipartite tomato-infecting Begomovirus species.     

Segregation distortion of Brassica carinata derived black rot resistance in Brassica oleracea

Three segregating F₂ populations were developed by self-pollinating 3 black rot resistant F₁ plants, derived from across between black rot resistant parent line 11B-1-12 and the susceptible cauliflower cultivar ‘Snow Ball’. Plants were wound inoculated using 4 isolates ofXanthomonas campestris pv. campestris (Xcc) race 4, and disease severity ratings of F₂ plants from the three populations were scored. A total of 860 arbitrary oligonucleotide primers were used to amplify DNA from black rot resistant and susceptible F₂ plants and bulks. Eight RAPD markers amplified fragments associated with completely disease free plants following black rot inoculation,which segregated in frequencies far lower than expected. Segregation of markers with black rot resistance indicates that a single, dominant major gene controls black rot resistance in these plants. Stability of this black rot resistance gene in populations derived from 11B-1-12 may complicate introgression into B. oleracea genotypes for hybrid production. This revised version was published online in July 2006 with corrections to the Cover Date.     

Diversity among Finnish net blotch isolates and resistance in barley

Summary Seedlings of a differential barley (Hordeum vulgare L.) series (21 genotypes) and of six check genotypes were used in the greenhouse to assess variation in virulence among 20 single-spore isolates of the net blotch pathogen. Pyrenophora teres Drechs. f.teres Smedeg., collected from various sites in Finland. The experiment was run twice and symptom expression was recorded on the first three leaves. Analysis of second leaf symptom scores from Run 1 indicated differences in virulence between isolates, all of which were pathogenic, and differential resistance among the barleys. The virulence of P. teres isolates appeared to be conditioned by the host barley from which the isolate derived; the average virulence of isolates collected from a susceptible host was greater than that of isolates collected from a less susceptible host. Results from Run 2 were similar regarding resistance within the barleys, but variation in virulence among the P. teres isolates was not consistent with that from Run 1. CI 9819 caries duplicate genes for resistance to P. teres (Rpt1b and Rpt1c), and CI 7548 possesses Rpt3d. Both genotypes were highly resistant to all isolates. The Rpt1a gene of Tifang (CI4407) conferred resistance to all isolates in Run 2, but only to about half of the isolates in Run 1. The checks, including two of the symptomatically most resistant Nordic barley genotypes, were universally susceptible during these stringent tests. No selective pressure has been placed on Finnish isolates of P. teres through previous deployment of major resistance genes, and it is speculated that any variation in virulence among isolates is likely to be due to a combination of evolutionary forces including, natural selection, random genetic drift and gene flow.     

Effect of leaf age on components of partial resistance in rice to leaf blast

Summary Ten rice genotypes were inoculated with a virulent isolate of Magnaporthe grisea to study the effect of leaf age on components of partial resistance, and evaluate associations between these components. The number of sporulating lesions per cm² leaf declined with increase of leaf age in all genotypes. The number of lesions per cm² leaf area in one week old leaves was about 25% that of the number in very young leaves in the susceptible cultivar CO39, but less than 2% in the more resistant cultivars IR36, IR60 and IR62. Large differences between genotypes were found for the number of sporulating lesions that developed, and this factor was closely related to the period that leaves remained susceptible after appearance. The number of lesions in the most susceptible cultivar CO39 was about 7 times that in the cultivars IR60 and IR64. Differences between genotypes were also found for lesion size. The effect of aging on average lesion size was less pronounced than on lesion density. Lesion size and lesion density were positively correlated, but a rapid decline of density was not necessarily accompanied by a rapid decline of size. No apparent differences between genotypes were observed for latent period. Genotypes with leaves that became highly resistant soon after appearance expressed higher levels of partial resistance in the field.     

Using molecular markers to pyramid genes for resistance to ascochyta blight and anthracnose in lentil (Lens culinaris Medik)

Ascochyta blight caused by the fungus Ascochyta lentis Vassilievsky and anthracnose caused by Colletotrichum truncatum [(Schwein.) Andrus & W.D. Moore] are the most destructive diseases of lentil in Canada. The diseases reduce both seed yield and seed quality. Previous studies demonstrated that two genes, ral1 and AbR1, confer resistance toA. lentis and a major gene controls the resistance to 95B36 isolate of C. truncatum. Molecular markers linked to each gene have been identified. The current study was conducted to pyramid the two genes for resistance to ascochyta blight and the gene for resistance to anthracnose into lentil breeding lines. A population (F_6:7) consisting of 156 recombinant inbred lines (RILs) was developed from across between ‘CDC Robin’ and a breeding line ‘964a-46’. The RILs were screened for reaction to two isolates (A1 and 3D2) ofA. lentis and one isolate (95B36) ofC. truncatum. χ² analysis of disease reactions demonstrated that the observed segregation ratios of resistant versus susceptible fit the two gene model for resistance to ascochyta blight and a single gene model for resistance to anthracnose. Using markers linked to ral1 (UBC 227₁₂₉₀), to AbR1(RB18₆₈₀) and to the major gene for resistance to anthracnose (OPO6₁₂₅₀),respectively, we confirmed that 11 RILs retained all the three resistance genes. More than 82% of the lines that had either or both RB18₆₈₀ and UBC227₁₂₉₀markers were resistant to 3D2 isolate and had a mean disease score lower than 2.5. By contrast, 80% of the lines that had none of the RAPD markers were susceptible and had a mean disease score of 5.8. For the case of A1 isolate of A. lentis, more than 74% of the lines that carriedUBC227₁₂₉₀ were resistant, whereas more than 79% of the lines that do not have the marker were susceptible. The analysis of the RILs usingOPO6₁₂₅₀ marker demonstrated that 11out of 72 resistant lines carried the marker, whereas 66 out of 84 susceptible lines had the marker present. Therefore, selecting materials with both markers for resistance to ascochyta blight and a marker for resistance to anthracnose can clearly make progress toward resistance in the population. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of a resistance specific to tomato spotted wilt tospovirus in Capsicum chinense ‘PI 159236’

Summary Inheritance studies were conducted to determine the genetic basis of resistance in pepper against one Tospovirus isolate classified as tomato spotted wilt virus (TSWV). F₁, backcrosses and F₂ populations were developed using the resistant parent Capsicum chinense PI 159236 (CNPH 679) and the susceptible parent C. annuum Magda (CNPH 192). Segregation ratios strongly indicated that the resistant response (a localization, hypersensitive-like reaction) to TSWV fits a single-dominant gene model. Under our experimental conditions, the penetrance of this gene was very high. This gene (tentatively named Tsw) is highly effective only against TSWV isolates. The resistance governed by the Tsw gene was not effective against isolates belonging to tomato chlorotic spot virus (TCSV) and groundnut ring spot virus (GRSV), two other previously described Tospovirus species.     

Inheritance of resistance to angular leaf spot in Nicotiana tabacum L. cultivars burley 21 and kentucky 14

Summary The genetics of resistance to angular leaf spot caused by Pseudomonas syringae pv. tabaci in Nicotiana tabacum cultivars Burley 21 and Kentucky 14 was investigated by studying disease reactions to three isolates of parental, F₁, F₂ and backcross generations derived from crosses between the resistant cultivars and the susceptible cultivar Judy's Pride. Studies were conducted in the greenhouse and in field plant beds. Chi-square values were computed to determine whether the observed ratios for disease reactions deviated from expected Mendelian ratios for a single, dominant gene controlling resistance to angular leaf spot in tobacco. Based on the resistance of the F₁ and the backcross generation to the resistant parent (BC-R), a 3 resistant: 1 susceptible segregation ratio in the F₂, and a 1 resistant: 1 susceptible segregation ratio in the backcross generation to the susceptible parent (BC-S), it was concluded that resistance to three isolates of Pseudomonas syringae pv. tabaci is governed by a single, dominant gene.     

A new gene for resistance in rice to Asian rice gall midge (Orseolia oryzae Wood Mason) biotype 1 population at Raipur, India

The Asian rice gall midge, Orseolia oryzae Wood Mason (Diptera: Cecidomyiidae), is a major pest of rice in several South and South East Asian countries. The maggots feed internally on the growing tips of the tillers and transform them into tubular galls, onion leaf-like structures called ‘silver shoots’ resulting into severe yield loss to the rice crop. We studied the mode of inheritance and allelic relationships of the resistance genes involved in resistant donor Line 9, a sib of a susceptible cultivar ‘Madhuri’. The segregation behaviour of F₁, F₂ and F₃ populations of the cross between Line 9 and susceptible cultivar MW10 confirmed the presence of a single dominant gene for resistance. Tests of allelism with all the known genes giving resistance to this population indicated that Line 9 possessed a new gene which was designated Gm 9 This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of reaction to Pseudomonas lachrymans in pickling cucumber

Summary Cucumber (Cucumis sativus) lines resistant to angular leafspot caused by Pseudomonas lachrymans react to an infection by developing necrotic lesions that lack the chlorotic halo characteristic of the susceptible reaction. The inheritance of the non-halo lesion reaction was studied in all possible crosses between resistant lines MSU 9402 and Gy 14A, and susceptible cultivars Wisc. SMR 18 and National Pickling. Genetic analysis of the F₁, F₂, backcross and F₃ populations revealed that the non-halo lesion type, associated with resistance, was controlled by a single recessive gene, pl. This character appears to be an important component of resistance to P. lachrymans.     

Response to selection for high and low partial resistance to leaf blast in F2 populations of three rice crosses

Summary Divergent selection for higher and lower partial resistance to leaf blast was applied in F₂ populations of crosses between the rice cultivars IR36, IR64 and CO39 after exposure to a virulent isolate. IR36 and IR64 are partially resistant while CO39 is highly susceptible. As selection criteria the number of sporulating lesions in leaves of the main culm or the lesion density in the topmost leaf relative to other F₂ plants with the same stage of development were used. A highly significant and meaningful response to selection was obtained in most cases, but the heritability was low. Realized heritabilities varied from 0.14 to 0.25 depending on the cross and were similar for both selection criteria. Selection for improved partial resistance to leaf blast is possible as early as the F₂. The efficiency of selection is probably much higher if replicated tests could be made, and better results are therefore expected if selection among F₃ lines is carried out. The results indicated that the relatively low infection efficiency in IR36 and IR64 is oligo- or polygenically controlled.     

Inheritance and allelism of resistances to the Russian wheat aphid in seven wheat lines

Summary Studies were conducted to determine the inheritance and allelic relationships of genes controlling resistance to the Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), in seven wheat germplasm lines previously identified as resistant to RWA. The seven resistant lines were crossed to a susceptible wheat cultivar Carson, and three resistant wheats, CORWA1, PI294994 and PI243781, lines carrying the resistance genes Dn4, Dn5 and Dn6, respectively. Seedlings of the parents, F₁ and F₂ were screened for RWA resistance in the greenhouse by artificial infestation. Seedling reactions were evaluated 21 to 28 days after the infestation using a 1 to 9 scale. All the F₁ hybrids had equal or near equal levels of resistance to the resistant parent indicating dominant gene control. Only two distinctive classes were present and no intermediate types were observed in the F₂ segregation suggesting major gene actions. The resistance in PI225262 was controlled by two dominant genes. Resistance in all other lines was controlled by a single dominant gene. KS92WGRC24 appeared to have the same resistance gene as PI243781 and STARS-9302W-sib had a common allele with PI294994. The other lines had genes different from the three known genes.     

Diallel analysis of resistance to Septoria tritici isolates in durum wheat

Summary All crosses, except for reciprocals, were made among ten cultivars originating from crop improvement programs in North Africa and the Middle East. The entries varied widely in reaction to Septoria tritici. F₁ and F₂ progenies of the crosses were evaluated using eight S. tritici isolates from seven countries in the Mediterranean area. Thus, sixteen separate combining ability analyses were excecuted. General combining ability (GCA) was the major component of variation, although specific combining ability (SCA) was present in most cases. Additive variance thus appears to be of predominant importance. Nevertheless, non-additive variance may interfere when line selection in a breeding program is practiced. While differing greatly among cultivars, specific GCA effects for each cultivar separately were of similar magnitude for all isolates. Ranking statistics determined that cultivars were ranked in similar order for both means and specific GCA effects independent of the isolate used. Different isolates may therefore interact with similar or identical genetically controlled mechanisms in a particular cultivar. This could indicate the absence of differential gene-for-gene relationships and suggests that isolates vary in aggressiveness rather than in virulence.     

Inheritance of resistance to the chlorosis component of tan spot of wheat caused by Pyrenophora tritici-repentis, races 1 and 3

The fungus Pyrenophora tritici-repentis, causal agent of tan spot of wheat, produces two phenotypically distinct symptoms, tan necrosis and extensive chlorosis. The inheritance of resistance to chlorosis induced by P. tritici-repentis races 1 and 3 was studied in crosses between common wheat resistant genotypes Erik, Hadden, Red Chief, Glenlea, and 86ISMN 2137 and susceptible genotype 6B-365. Plants were inoculated under controlled environmental conditions at the two-leaf stage and disease rating was based on presence or absence of chlorosis. In all the resistant × susceptible crosses, F₁ plants were resistant and the segregation of the F₂ generation and F₃ families indicated that a single dominant gene controlled resistance. Lack of segregation in a partial diallel series of crosses among the resistant genotypes tested with race 3␣indicated that the resistant genotypes possessed␣the same resistance gene. This resistance gene was effective against chlorosis induced by P.␣tritici-repentis races 1 and 3.     

Resistance to Septoria nodorum blotch in several Triticum species

Summary Resistance to septoria nodorum blotch in Triticum monococcum, T. tauschii, T. timopheevii, T. dicoccum and T. durum was evaluated on plants at the three-leaf stage in greenhouse tests. A high frequency of resistant genotypes was found in T. monococcum, T. tauschii and T. timopheevii, but not in T. dicoccum and T. durum. The resistance of F₁ plants of crosses of resistant T. monococcum (PI 289599) and T. timopheevii (PI 290518) accessions with susceptible common wheat cv. Park and durum wheat cv. Wakooma, respectively, was evaluated on the basis of percentage leaf necrosis, lesion number, lesion size and incubation period. No dominance was found for long incubation period, but various dominance relationships occurred for low percentage leaf necrosis, low lesion number and small lesion size, depending on the cross. Multiple regression analysis showed that lesion number contributed more to percentage leaf necrosis than lesion size or incubation period. Resistance to septoria nodorum blotch was transferred successfully from T. timopheevii to cultivated durum wheat. Resistant BC₁F₇ lines, recovered from the T. timopheevii (PI 290518) × Wakooma cross, showed normal chromosome behaviour at meiosis (14 bivalents) and were self-fertile. However, an effective level of resistance was not recovered in lines derived from the other interspecific crosses.     

A new powdery mildew resistance gene: Introgression from wild emmer into common wheat and RFLP-based mapping

An Israeli accession (TTD140) of wild emmer, Triticum turgidum var. dicoccoides, was found resistant to several races of powdery mildew. Inoculation of the chromosome-arm substitution lines (CASLs) of TTD140, in the background of the Israeli common wheat cultivar ‘Bethlehem’ (BL), with five isolates of powdery mildew revealed that only the line carrying the short arm of chromosome 2B of wild emmer (CASL 2BS) exhibited complete resistance to four of the five isolates. To map and tag the powdery mildew resistance gene, 41 recombinant substitution lines, derived from a cross between BL and CASL 2BS, were used to construct a linkage map at the gene region. The map, which encompasses 69.5 cM of the distal region of chromosome arm 2BS, contains six RFLP markers, a morphological marker (glaucousness inhibitor, W1 ^I), and the powdery mildew resistance gene. Segregation ratios for resistance in F₂ of BL × CASL 2BS and in the recombinant lines, combined with the susceptability of F₁ progeny to all tested isolates, indicate that resistance is controlled by a single recessive allele. This alleleco-segregated with a polymorphic locus detected by the DNA marker Xwg516, 49.4 cM from the terminal marker Xcdo456. The new powdery mildew resistance gene was designated Pm26. This revised version was published online in July 2006 with corrections to the Cover Date.     

Variation in virulence and resistance in the bean-bean rust pathosystem in Ecuador

Twenty two Uromyces appendiculatus isolates were tested on 20 differential and 25 Ecuadorian Phaseolus vulgaris cultivars in the seedling stage. Based on the infection types 20 races could be discerned. The Ecuadorian cultivars differed greatly in their reaction to the isolates, from resistant to only one isolate (`Red Small Garden') to resistant to all isolates (’G2333‘). The isolates showed a wide range of virulence to the Ecuadorian cultivars, from virulent to only two cultivars (isolate 13) to virulent to 21 cultivars (isolates 5 and 23). Seven cultivars with a basically susceptible infection type appeared to differ greatly in quantitative resistance when tested in three consecutive crop cycles. The disease severities in percentage leaf area affected averaged over the three cycles ranged between 83.9% for ‘Red Small Garden’ and 13.1% for ‘INIAP-414’. Race-specific resistance does not seem an advisable breeding strategy, but the quantitative resistance offers a good alternative.