Inheritance of resistance to two Septoria tritici isolates in spring and winter bread wheat cultivars

Summary All possible crosses (including reciprocals) were made among four winter bread (Aurora, Bezostaya 1, Kavkaz, and Trakia) and two Israeli spring wheat cultivars (spring x winter diallel), and among two South American spring wheats (Colotana and Klein Titan) with the same Israeli cultivars (spring x spring diallel) to study the inheritance of resistance to septoria tritici blotch. Parents, F₁, F₂ and backcrosses were grown in two separated blocks in the field over two years. One block was inoculated with isolate ISR398A1 and another with ISR8036. Each plant was assessed for plant height (cm), days to heading (from emergence or transplanting), and percent pycnidia coverage on the four uppermost leaves. Plant height and maturity had insignificant effects on pycnidia coverage. No cytoplasmic effects could be detected. In the spring x winter diallel general combining ability (GCA) was the major component of variation. Significant specific combining ability (SCA) was present in all cases. Partial dominance was operative in populations inoculated with ISR398A1. Resistance in the winter wheats was controlled by a small number of genes (usually two). The four winter wheats derive their resistance to ISR398A1 from their common parent Bezostaya 1 which lacks the 1B/1R wheat-rye translocation. Their resistance is readily overcome by ISR8036. Inheritance of the South American wheats can be explained by additive effects, with a small number of genes of recessive mode affecting resistance to both isolates. Breeding strategies that favor additive, and additive x dominance gene action should be pursued.     

Variations in pathogenicity patterns ofMycosphaerella graminicola withinTriticum spp. in Israel

Summary Pathogenicity patterns of 42Mycosphaerella graminicola (Septoria tritici) isolates secured from the major wheat growing regions in Israel, were assessed on seedlings of 16 bread and durum wheats. The spring bread wheat cultivar Titan (CI12615), the winter bread wheats Bezostaya 1, Kavkaz, NE7060 and the durum wheats Nursit 163 and Zenati Bouteille, all exhibited a high level of resistance. Significant cultivar × isolate interactions were recorded for isolates from the different regions in relation to the response of the 16 cultivars organized in 6 reponse classes to the test isolate ISR8036. Number of genes for resistance in the 16 cultivars was estimated, based on the assumption of a gene-for-gene relationship. Twelve complementary genes were hypothesized in the 42M. graminicola isolates × 16 wheat cultivars matrix. There was considerable variation in virulence frequencies between regions and between locations within the same region. The overall frequency of Bezostaya-Kavkaz virulence was low throughout the country (5–7%). The frequency of designated hypothesized virulence genes VST OLAF, VST COLOTANA, VST IAS 20 in Israel, was high in most locations. The reported analytical approach enables the identification of virulence hot-spots and assists in designing breeding for resistance strategies.     

The combined effect of plant stature and maturity on the response of wheat and triticale accessions to Septoria tritici

Summary The relationships between percent pycnidia coverage on the four uppermost leaves (PCD), plant height (PHT) and days to heading (HED) were evaluated for 21,000 wheat and triticale accessions tested in artificially inoculated (with fixed combination of S. tritici isolates) field nurseries over 8 trial years. A general Linear Model procedure (GLM) estimated Septoria severity using two correlative models: model 133-1 Year and model II–PCD=b₁PHT+b₂HED+C. The regression coefficients for PHT and HED in the two models were –0.54 and –0.40, respectively, with a R²=0.80^** and R²=0.29^** for model I and model II, respectively. The predicted cultivar best fitted to the model would be characterized as a semidwarf (PHT=115 cm) with an early-moderate maturity (HED=95 days to heading). The estimated mean percent pycnidial coverage for the two models over the 8 trial years was 40.8%. The performance of a group of 38 cultivars replicated yearly during the 8 trial years was assessed relative to model I. The deviation of each cultivar from the model was calculated using two functions: a) Sum Relative Serial Deviation (SRSD) and b) Total Relative deviation (TRD), in addition to Standard errors (SE). The proposed analytical protocol enabled identification of cultivars which expressed consistent yearly deviation (from the model) in host response combined with low-moderate mean pycnidial coverage (±30%). Such cultivars may possess a more stable type of genetic protection against the adverse effects of septoria tritici blotch.     

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.     

Evaluation of screening techniques and determination of criteria for assessing resistance to Corynebacterium michiganense in tomato

Summary When screening populations segregating for resistance to Corynebacterium michiganense a rapid and reliable inoculation technique is needed. From the several techniques tested, inoculation of the petiole was found to give better differentiation of resistant and susceptible plants than did inoculation of the root or leaf. When petiole-inoculation was compared with stem-inoculation the latter showed the lowest environmental variation and is suggested for use in genetic studies. Stem-inoculation involved clipping the stems of 4 week old seedlings 1 cm above the cotyledonary leaves with scissors which had been dipped in inoculum.Petiole inoculation and, to a lesser degree, stem inoculation failed to produce uniform infection, resulting in variation in the expression of the disease. The scoring was therefore based on the presence, and not the degree, of wilting and stunting, or the size of the cankers. Plants free of all these symptoms were rated resistant. The coefficient of variation for vascular discoloration on individual plants was calculated to be 50%, which in part may be explained by the histopathology of the host.The virulent isolates cm 21 caused death of seedlings resistant to isolate H. However, cultivars resistant to isolate H in the seedling stage did not decrease in fruit weight as compared to susceptible cultivars when inoculated with isolate cm 21 at anthesis.Michigan Agricultural Experiment Station Journal No. 7424.     

Pathotypes and NIP1 gene sequences of Finnish Rhynchosporium secalis isolates from barley, couch grass and rye

Rhynchosporium secalis (Oud.) J.J. Davis, causal agent of scald of barley (Hordeum vulgare L.), was isolated from barley, couch grass (Elymus repens L.) and rye (Secale cereale L.). Isolates were used to inoculate seedlings of a differential barley series containing several sources of major gene resistance to the disease. The series included Atlas46, (resistance gene Rrs1) and the isogenic line Atlas, which lacks the gene (rrs1). The necrosis-inducing peptide NIP1 has been suggested to be the product of the avirulence gene AvrRrs1 (NIP1) that with the barley resistance gene Rrs1 determines the incompatibility of the pathogen-host interaction. All R. secalis isolates were virulent only on the susceptible barley cultivars Arve and Chan, irrespective of spore concentration or original host species. There were no indications of redundant virulence among the R. secalis isolates. The NIP1 gene was sequenced from each isolate and there was no correlation with source of the isolate or sequence modification and virulence. Four isolates, from barley and couch grass, were characterised by a basic Type I NIP1sequence. The recorded NIP1 sequence changes are consistent withR. secalis populations not receiving selection through deployment of the Rrs1 gene in commercial barley cultivars in Finland. This revised version was published online in July 2006 with corrections to the Cover Date.     

Residual effects of the Xa-4 resistance gene in three rice cultivars when exposed to a virulent isolate of Xanthomonas campestris pv. oryzae

Summary F₂ plants of five, and F₃ plants of three, crosses between genotypes carrying the race-specific resistance gene Xa-4 and genotypes not carrying this gene were inoculated with two isolates of Xanthomonas campestris pv. oryzae. Half the tillers of each plant received isolate PX061, avirulent on the Xa-4 gene, the other half of the tillers received isolate PX099, virulent for the Xa-4 gene. The F₂ and F₃ populations segregated for a single dominant resistance gene, Xa-4.The parental, F₂ and F₃ genotypes not carrying Xa-4 had mean lesion lengths between 28 and 29 cm for both isolates. The Xa-4 carrying parents showed a mean lesion length of 2.7 cm with the avirulent isolate and of 12.4 cm with the virulent isolate. The Xa-4 carrying F₂ and F₃ genotypes had mean lesion lengths of 5.2 and 20.1 cm for the two isolates, respectively. These observations strongly indicate that the Xa-4 gene, carried by the rice genotypes studied (IR28, Cisadane and BR51-282-8), had a considerable residual effect when exposed to virulent isolate PXO99.     

Leaf scorch (Kabatiella caulivora (Kirch.) Kirk.) of subterranean clover (Trifolium subterraneum L.): Influence of host cultivar,growth stage and pathogen isolates on the disease severity

Summary Interaction of six isolates of Kabatiella caulivora (545, 547, 2212, BNL, BDG and KLN) collected from different geographical regions of Western Australia and Victoria with six cultivars (Clare, Daliak, Enfield, Guildford D, Karridale and Woogenellup) of subterranean clover was studied at seedling (2-week old) and adult-plant (12-week old) stages. Seedling response measured as percentage of diseased plants showed cultivar Daliak to be immune to all isolates, while the remaining cultivar x isolate combinations showed differing degrees of susceptibility.The adult-plant response, measured as percentage of petioles affected per plant, showed that all the cultivars were susceptible to varying degrees to all the isolates, with Daliak being the most resistant cultivar.Analysis of variance of both seedling and adult-plant resistance showed significant (P<0.05) effects due to cultivars, isolates and their interaction. The magnitude of variance due to the isolates was substantially reduced when measured as adult-plant response. Leaf scorch severity was clearly influenced by cultivar and its developmental stage, and by isolate of the pathogen.     

Response of wheat cultivars to infection by Stagonospora nodorum isolates/mixture on detached and intact seedling leaves

Stagonospora nodorum blotch (SNB) of wheat caused by Stagonospora nodorum (Berk.) Castellani & E.G. Germano is among the major foliar diseases of wheat worldwide. The response of four wheat cultivars for reaction to infection by isolates/mixture of S. nodorum was tested on detached and intact seedling leaves in the laboratory and glasshouse experiments, respectively. The four wheat cultivars tested significantly differed from each other in response to infection by the isolates/mixture of S. nodorum. Similarly, the aggressiveness of the four isolates/mixture of the pathogen on the four wheat cultivars was significantly different as demonstrated by the parameters tested in the two experiments. However, no significant (p ≤ 0.05) isolate x cultivar interaction was observed for all the parameters tested in the two experiments. Highly significant positive or negative correlations were observed between parameters, i.e., disease severity (DS), incubation period(IP), and size of necrotic lesions (SNL), tested on detached and intact seedling leaves, respectively. This suggests that use of the detached leaf technique has considerable promise for quick screening of wheat cultivars against SNB. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Stable resistance in barley to Pyrenophora teres f. teres isolates from the Nordic-Baltic region after increase on standard host genotypes

Results from tests of a mixture of Finnish net blotch, Pyrenophora teres Drechs. f. teres Smedeg., isolates on a differential series of barley seedlings, comprising 17genotypes, indicated that patterns of infection response (IR)and percentage leaf area damaged (PLAD) were unaffected by differences in seedling size. Variation of the concentration of inoculum between 1,250 conidia ml^-1 and 20,000 conidia ml^-1 produced similar patterns of IR and PLAD on the differential series. IR and PLAD scored on the second seedling leaf differentiated resistance to P. teres f. teresamong the genotypes better than on the first seedling leaf. Ina second experiment, 120 single-spore P. teres f. teres isolates from Finland, Sweden, Norway, Latvia, Estonia and Ireland were used in tests conducted in the greenhouse to differentiate them in terms of virulence reaction on seedlings of six differential barley genotypes. Each isolate was tested directly following isolation from the leaf material and after having passaged each through barley cvs. Arve or Pohto, to produce 360 isolates in total. Virulence of the isolates differed significantly on the members of the differential series, but differences associated with country of origin and passaging, and interactions, were small. It is concluded that little variation between virulence of P. teres f. teres isolates is evident over a large geographic area, incorporating Nordic and Baltic countries, and Ireland. Barley genotype response to P. teres f. teres appeared to be of more significance than relative virulence of the pathogen isolates. This could simplify breeding barley for improved resistance to this phytopathogen. This revised version was published online in August 2006 with corrections to the Cover Date.     

Characterization of the resistance to <Emphasis Type="Italic">Phytophthora infestans</Emphasis> in local potato cultivars in Bolivia

This experiment was carried out to investigate whether and how much field resistance to late blight, caused by Phytophthora infestans, is present in the local cultivated potato germplasm. In total 36 entries were compared in a field experiment in an area highly conducive to late blight development. Of the 36 cultivars 32 were local cultivars belonging to five Solanum species, S. tuberosum (1 accession), S. andigena (18), S. juzepczukii (2), S. stenotomum (9) and S. ajanhuiri (2). The other four cultivars were derived from breeding programmes, one being the Dutch cultivar Alpha used as a highly susceptible control. The 36 cultivars were planted according to a simple 6 × 6 lattice design with three replicates. Each replicate was divided in six incomplete blocks each with six cultivars. The disease severity was assessed weekly during 9 weeks starting 48 days after planting. The area under the disease progress curve (AUDPC) was used as a measure of the field resistance. Nine isolates from surrounding potato fields were tested for their virulence to the resistance genes R1–R11 using 22 differential cultivars. The components of the field resistance of 19 of these cultivars were compared in the greenhouse using a local isolate with virulence to all known R-genes, except to R9. The nine isolates represented seven races with a race complexity varying from 7 to 10 virulence factors. All isolates carried virulence against R1, R2, R3, R7, R10 and R11, while virulence against R9 was absent. The AUDPC among the 32 local cultivars ranged from very large, significantly larger than that of ‘Alpha’ to very small. The AUDPC from S. stenotomum accessions ranged from very large to intermediate, those from S. andigena accessions from large to very small. Especially among the S. andigena accessions interesting levels of field resistance were found. Four components of field resistance were assessed, latency period (LP), lesion size (LS), lesion growth rate (LGR) and relative sporulation area (RSA). All four showed a considerable variation among the cultivars. The LP ranged from 3½ to 6 days. The LS ranged from 225 mm² to 20 mm². The LGR varied about six-fold, the RSA more than 10-fold. The components tended to vary in association with one another. LP and LGR were well associated with each other and had a significant correlation with the AUDPC.     

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.     

Assessment of quantitative resistance of rice cultivars to Xanthomonas campestris pv. Oryzae: a comparison of inoculation methods

Summary Lesion size and lesion number were measured on cultivars of rice inoculated by clipping or spraying with virulent isolates of Xanthomonas campestris pv. oryzae. Mean percentage diseased leaf area (%DLA) gave a similar ranking for the two inoculation methods but differences in lesion size among cultivars were much more evident after clip than after spray inoculation. Correlation between the methods was high (r=0.82^**), but some cultivars responded differently with the two techniques. Cultivars which had low scores following spray inoculation showed low disease progress during the first nine weeks after transplanting into a screen-house experiment. Assessment after clip inoculation measures resistance due to spread of bacteria within the leaf xylem system, an important component of quantitative resistance. Assessment after spray inoculation measures all resistance, including resistance to entrance of bacteria into the leaf. In order to select rice entries with improved quantitative resistance to X. c. oryzae based on both components, a screening based on lesion length after clip inoculation, followed by a test for lesion number after spray inoculation, is advised.     

Race differentiation and search for sources of resistance to Rhynchosporium secalis in barley in Italy

Summary Barley in Italy has recently been seriously affected by Rhynchosporium secalis. The pathogenic variation of the fungus was studied and 17 races were differentiated on 13 barley cultivars carrying most of the currently known genes for resistance. RC 1, the most virulent and most frequent race, was virulent on 10 out of the 13 differentials and the remaining races proved to be less virulent variants of RC 1. Atlas (C.I. 4118), Atlas 46 (C.I. 7323) and Osiris (C.I. 1622) were the only three differentials resistant to all the analyzed single-spore isolates.Differential cultivars previously assumed to have identical resistance factors did not react in the same way to all the Italian races, thereby revealing either undisclosed differences in the genes described or the presence of additional unidentified ones.Our findings were compared with previous data about virulence of scald populations from different countries, on the basis of tests with common differentials: fundamental differences were found between the Italian population and those of other countries with regard to virulence patterns.The susceptible reactions to race RC 1 of most barley cultivars grown in Italy indicate the urgent need for resistance genes to be incorporated in the cultivated material. Seventy-one barley accessions, known as sources of resistance in different parts of the world, were screened for their behaviour to races RC 1 and RC 13. Twenty-two appeared resistant to both of them.     

Pathogenicity of the Rhynchosporium secalis population in the Western Cape province of South Africa

The virulence spectra of 50 Rhynchosporium secalis isolates from a population in the Western Cape province of South Africa were determined, and 21 races were detected when evaluated against 17 differential cultivars. The virulence spectrum of the R. secalis population shows considerable variation, and carries unnecessary virulence genes which is quite unexpected, since chiefly susceptible barley cultivars are grown in the south Western Cape. The two most prevalent races, namely races 4 and 7 had three and four virulence genes respectively. Both race 4 and 7 were virulent on the most susceptible cultivars, West China, Steudelli, C.I.8618 and C.I.2226. Considering the resistance genes reported for the cultivars Atlas 46, Turk, and C.I.3515 which showed no susceptible cultivar-pathogen interaction, it would appear that the Rh-Rh3-Rh4 complex is primarily involved in conferring resistance to the local R. secalis isolates This revised version was published online in July 2006 with corrections to the Cover Date.     

Rust infections of some Allium species: An assessment of germplasm for utilizable rust resistance

Summary Isolates of three species of rust (Puccinia allii, P. mixta and Uromyces ambiguus) were each inoculated onto a range of potential host plants, selected to include the major European crop Allium species together with the commoner wild species in the section Allium. Measurements of incubation period, lesion type, latent period and pustule quantity were made in a single environment. Each isolate was able to sporulate on a range of Allium species. There was evidence of specialisation to particular sections of the genus; all the leek isolates performed well on the majority of hosts in section Allium, whilst the chive isolate grew poorly on most members of that section. Exceptions to this are exemplified by A. fistulosum being attacked by all three rust species and A. sativum being attacked by both leek and chive isolates. A continuum of reaction types was found which, for the leek isolates, ranged from the low levels of quantitative resistance in leek cultivars, through higher quantitative levels within the A. ampeloprasum complex, to qualitative resistance in several interactions in non-host sections of Allium, where the pathogen was able to infect and form colonies of some size without sporulating.     

Evaluation of the virulence of races 1, 2 and 4 of Fusarium oxysporum f.sp. dianthi in carnation

Summary A simultaneous analysis of the virulence of races 1, 2 and 4 of Fusarium oxysporum f.sp. dianthi to a series of nine carnation cultivars revealed the presence of different interactions between races and cultivars, as well as differences in pathogenesis between race 1 on the one hand and race 2 and 4 on the other.The most common race 2 induced typical symptoms of Fusarium wilt in all susceptible cultivars. The cultivars showed considerable variation in resistance to race 2. Only Novada remained free of external symptoms throughout the experiment. In diseased plants of all cultivars studied, infected vascular tissue was white with dark brown margins, and heavy degradation of the cell walls and xylem parenchyma cells had occurred. All Dutch isolates corresponded with race 2.Race 4 induced wilt symptoms similar to those induced by race 2, and there was a similar variation in resistance to race 2 and 4 in the cultivars. On average, the race 4 isolates were less aggressive than those of race 2. Compared with race 2, there was evidence of some genotype × race interactions: Pallas proved to be considerably more susceptible, and Lena more resistant to race 4 than to race 2. The isolates of race 4 induced a nistopathology similar to that induced by race 2, but with less vascular browning.Race 1 induced atypical but severe wilt symptoms and unusual vascular discoloration in Elsy, Niky and Sam's Pride only. The vascular tissue in these cultivars turned pale brown; in spite of heavy colonization of these tissues virtually no degradation of cell walls was observed. All other cultivars tested proved virtually resistant to race 1, providing further evidence for genotype × race interactions.Within races, limited but statistically significant genotype × isolate interactions were found as well, in particular within race 4. These are tentatively attributed to independent variation of two (or more) resistance components.     

Genes for resistance in barley to Finnish isolates of Rhynchosporium secalis

Summary Twenty Finnish isolates of Rhynchosporium secalis (Oud.) J.J. Davis, the causal agent of scald, were taken from infected barley (Hordeum vulgare L.) plants and inoculated on to seedlings of a differential series of barley containing a range of major genes for resistance to the fungus, as well as on to six Nordic 6-row spring barleys and three winter ryes (Secale cereale L.). These fungal isolates derived from four sites and three host varieties. Disease development was monitored on two leaves of seedlings in the greenhouse employing a standard scale, and on adult plants in the field by assessing the diseased area on the three uppermost leaves. A comparison was also made between the pathogenicity and virulence of ten Finnish and ten Canadian R. secalis isolates. The Finnish isolates varied in virulence, but with the exception of Algerian (CI 1179) seedlings and adult La Mesita (CI 7565) all seedlings and adult plants of the entire differential series were resistant to all isolates. Canadian isolates were, on average, less virulent than Finnish isolates. All the Nordic checks were susceptible to all Finnish and seven Canadian isolates, but differences in the degree of susceptibility were evident. Isolates of R. secalis from barley were non-pathogenic on rye, isolates from Elymus repens L. were non-pathogenic on barley and rye, and isolates from rye were only pathogenic on rye. Finnish R. secalis isolates contain no redundant pathogenic diversity. The differential series represents a useful, but as yet untapped, source of resistance to R. secalis for Finnish barley breeders.     

Diallel analysis of four resistance components to Septoria tritici in six crosses of wheat (Triticum aestivum)

Four resistance components, i.e. incubation period, latent period, maturation period and pycnidial coverage, were studied in a diallel cross of four wheat cultivars differing in susceptibility to Septoria tritici. Inoculations and evaluations were carried out at the fourth leaf stage for two years. A combined ANOVA for both years exhibited a significant interaction between crosses and years for maturation period. A separate ANOVA for each year was done for this trait. The combined ANOVA showed that crosses were significantly different for incubation, latent period and pycnidial coverage. Cross effects were also significant for maturation period in each year for the separate ANOVA. Year effects were significant for latent period and maturation period. General combining ability (GCA) was significant and preponderant for incubation period, latent period and pycnidial coverage. For maturation period, the separated ANOVA showed the significance of the GCA for both years. Specific combining ability (SCA) was significant for incubation and pycnidial coverage and for maturation in one year. The preponderance of the additive genetic variance for all traits indicates the possibility of selecting for them in order to obtain improved cultivars. Incubation period was inherited independently of maturation period and pycnidial coverage, indicating that combinations of some of those characters may lead to more effective and durable resistance.