Inheritance of components of partial resistance of barley to Rhynchosporium secalis with particular reference to race specificity

Six spring barley cultivars with no known genes for resistance to specific virulences but varying in partial resistance to Rhynchosporium secalis, were crossed in all combinations (6 × 6 diallel including reciprocals). In addition to seeds from naturally selfed plants, seeds of all parent cultivars were also produced by artificial selfing (emasculation followed by pollination using pollen from the same cultivar). This ensured comparability between seeds of parents and F₁. Both sets of parents, F₁ and F₂ families were grown in the field as single spaced plants and inoculated at Zadoks growth stage 49 with spore suspensions (2 × 10⁶ spores ml^-1) of three races (pathotypes) of R. secalis (Zadoks, Chang & Konzak, 1974). Components of partial resistance, incubation period (ICP), infection frequency (IF) and spore production per lesion (SP/L) were assessed on each plant. There were highly significant differences for all three components of partial resistance in both sets of parent cultivars but rank order in both sets was similar as evidenced by correlation coefficients, r= 0.96 for ICP and IF and r= 0.87 for SP/L. All three components of partial resistance were strongly correlated with NIAB (National Institute of Agricultural Botany, Cambridge, UK) resistance ratings. Means of F₁ and F₂ families were correlated with mid-parent values for ICP and IF but not SP/L. No difference in aggressiveness was found between races but for each component of partial resistance there was a significant interaction between race and parent cultivar (artificial selfs) and, for IF and ICP, a significant interaction between race and F₁ family. There was no evidence of interaction between parent (natural selfs) and race nor between race and F₂ family. Examination of genetic control of resistance showed evidence of strong additive effects (combining ability) in both F₁ and F₂ for ICP and IF but not for SP/L. There was no evidence for maternal or reciprocal differences, but there was evidence for dominance effects although their nature differed between components of partial resistance and between F₁ and F₂ generations. In the F₁, but not the F₂ generation, several elements of dominance (direction, distribution of dominant genes between parent cultivars, specific combining ability) showed for ICP or IF (but not SP/L) significant interaction with race.     

Components of partial resistance as criteria for identifying resistance

The components of partial resistance, incubation period, lesion area, latent period and sporulation were recorded on plants of six winter and two spring wheat cultivars which had been artificially inoculated with Septoria nodorum spores. Incubation period gave a guide as to how the cultivars would respond in the field to Septoria nodorum but statistical analysis showed that it could not be used alone to predict accurately the resistance of each cultivar to the pathogen. Average sporulation, however, could be used with more confidence for predicting the field resistance of the cultivars. From a regression analysis of NIAB rating versus incubation period, lesion area, latent period and sporulation, an equation was devised to obtain resistance indices for each cultivar. These resistance indices clearly reflected the NIAB ratings for the cultivars. It would therefore appear that resistance indices could be used as a pre-field evaluation method for identifying resistance to Septoria nodorum and thus be a valuable technique in breeding programmes.     

The Expression of Partial Resistance of Wheat to Puccinia recondita

A range of winter wheat cultivars, which appeared to show no race-specific resistance to Puccinia recondita f. sp. tritici, was studied at seedling and adult growth stages to determine the components of partial resistance which might contribute to slow-rusting in the field. Under controlled environmental conditions, adult plants of cvs Atou, Bouquet, Cappelle-Desprez and Maris Freeman produced less uredospores per unit area than the control cv. Armada, and uredosori were smaller on these cultivars. Fewer uredosori erupted on cv. Cappelle-Desprez than on the control, and both cv. Bouquet and cv. Cappelle-Desprez showed longer latent periods than other cultivars. Cultivar Holdfast had a shorter latent period, with larger and more numerous uredosori than even the highly susceptible cv. Armada. Cultivar Waggoner displayed a similar response to infection to that of the control. The components of partial resistance detected in adult plants were not displayed by seedling leaves.     

Determination of Rust Resistance Gene Complex Lr34/Yr18 in Spring Wheat and its Effect on Components of Partial Resistance

The non‐durable nature of hypersensitive (race‐specific) resistance has stimulated scientists to search for other options such as race‐non‐specific resistance to provide long‐lasting protection against plant diseases. Adult plant resistance gene complex Lr34/Yr18 confers a dual race‐non‐specific type of resistance to wheat against stripe rust (Puccinia striiformis f. sp. tritici) and leaf rust (P. triticina Eriks). This study was conducted to evaluate 59 spring bread wheat (Triticum aestivum L.) genotypes for the presence of the Lr34/Yr18‐linked csLV34 allele using STS marker csLV34 and to determine the effect of this gene complex on the components of partial resistance in wheat to leaf/stripe rust. Lr34/Yr18‐linked csLV34 allele was detected only in 12 genotypes, namely Iqbal 2000, NR‐281, NR 354, NR 363, NR 364, NR 366, NR 367, NR 370, NR 376, 4^thEBWYT 509, 4^thEBWYT 510 and 4^thEBWYT 518. Eleven genotypes showing the amplified Lr34/Yr18‐linked allele were further studied for the assessment of the effect of Lr34/Yr18 on components of partial resistance along with nine genotypes lacking this gene complex. Both stripe and leaf rusts were studied separately. The components of partial resistance including latency period (LP) and infection frequency (IF) were studied on primary leaf (seedling stage), fourth leaf and fully expanded young flag leaf (adult plant stage). Both the stripe and leaf rust fungi showed a prolonged LP and reduced IF on genotypes carrying Lr34/Yr18 gene complex. Generally, a longer LP was associated with a reduced IF at all growth stages. Although significant effect of Lr34/Yr18 gene complex on LP and IF was observed almost at all three growth stages, the effect was more pronounced at flag leaf. This suggested that Lr34/Yr18 gene complex is more effective at later stages of plant growth.     

Levels and components of resistance to Amphorophora idaei in raspberry cultivars containing different resistance genes

Levels and components of resistance to Amphorophora idaei in raspberry cultivars containing different A. idaei resistance genes were studied under infestation tunnel, glasshouse and laboratory conditions. Each test consistently ranked raspberry cultivars, with increasing levels of resistance in the order, non-resistant (cv. Mailing Jewel), resistant cultivars containing minor genes, major gene A₁, gene A₁₀ (red raspberry) and gene A₁₀ (black raspberry) respectively. Resistance was expressed in three different ways; decreased alatae settling and feeding, decreased apterae settling and decreased aphid fecundity and rate of nymphal development. Following exposure to a large population of alatae, significantly fewer aphids settled on A₁, A₁₀ and to a lesser extent, minor gene-containing cultivars, compared to cv. Mailing Jewel. More alatae settled on the top than the bottom zone of non-resistant, minor gene resistant and two of four A₁-containing cultivars; alatae settling was low on all zones of A₁₀-containing raspberries. Aphid fecundity and nymph development patterns on different cultivars and resistance classes were similar to those found for alatae settling. After 7 days reproduction more than 30% of the nymphs developed to third or fourth instar on cv. Mailing Jewel, whilst on minor gene and major gene-containing cultivars the total number of nymphs and the proportion of later instars decreased. On resistant cultivars the nymphs were found mainly on the middle and bottom leaf zones, compared to cv. Mailing Jewel. A rapid (48 h) screening test using floating leaflets was developed and, on the basis of apterae settling, ranked cultivars reliably; it was particularly effective in distinguishing moderate and strong resistance to strain 1 A. idaei. Gene A₁-containing cultivars bred in England were much more resistant to strain 1 A. idaei than those bred in Scotland. Possible reasons for this difference are discussed, in relation to modified screening procedures and to the control of the viruses transmitted by this aphid vector.     

Composition of root exometabolites of the symbiotically effective pea cultivar triumph and its parental forms

The qualitative and quantitative composition of low-molecular exometabolites in roots of pea (Pisum sativum L.) was studied with a cultivar Triumph and its parental forms (a symbiotically effective variety k-8274 and a modern highly productive cv. Classic). A relationship between root exudation and the ability of cultivars to establish symbiosis was analyzed. In the early stages of plant growth, the roots of cv. Triumph exhibited low exudation of organic acids, sugars, and amino acids. The quantitative composition of organic acids in the root exudates of cv. Triumph was close to that of cv. k-8274, whereas the composition of sugars and amino acids was similar to that of cv. Classic. In the field experiment, the effect of inoculation with a mixture of rhizobium strains and mycorrhizal fungus on plant growth was more evident in cv. Triumph than in cvs. Classic and k-8274. The results suggest that the high symbiotic potential of cv. Triumph is related to exudation of pyruvic and succinic acids that were the major components of root exometabolites both in Triumph and k-8274 cultivars.     

Determining the effectiveness of resistance to subterranean clover mottle sobemovirus in different genotypes of subterranean clover in the field using the grazing animal as virus vector

The effectiveness of resistance to subterranean clover mottle sobemovirus (SCMoV) previously identified in different genotypes of subterranean clover (Trifolium subterraneum) inoculated with infective sap in the glasshouse, was tested in two field experiments which used the grazing animal as virus vector. Replicated plots each consisting of paired test rows of 20 different genotypes were used. Clover plants infected with SCMoV were transplanted in between the paired test rows and these acted as sources of the virus for spread by grazing sheep. Although used in different years at different sites with different virus isolates, the field exposure methodology employed produced consistent results. The genotypes each behaved similarly in both experiments as regards the relative extents of SCMoV infection that developed, levels ranging from 0–98%. The previously identified resistance in six ‘highly resistant’ and three ‘partially resistant’ cultivars was effective under field conditions. However, the ‘partial resistance’ in three others was overcome, cvs Green Range and Mt Barker developing levels of infection approaching those in ‘susceptible’ cultivars, while an intermediate infection level developed in cv. Karridale. The three cultivars in which partial resistance was not effective all belonged to ssp. subterraneum. In subterranean clover breeding programmes, field screening using the grazing animal as a vector is advisable to determine whether SCMoV resistance found by sap inoculation is still effective under field conditions.     

The inheritance of resistance to Rhynchosporium secalis in some European spring barley cultivars

The inheritance of partial resistance to Rhynchosporium secalis, which appears to be race non-specific, was studied in a diverse range of European spring barley cultivars. Data from the F₂ generation of a 6 × 6 diallel cross and the F₃ generation of three crosses selected from this diallel set suggested that resistance was complex in inheritance, the results being incompatible with any hypothesis involving less than four genes. The F₂ studies indicated that both dominant and recessive genes were active in conferring resistance, and that there were significant additive gene effects. Transgressive segregation occurred in all cross combinations in the F₃ material. Consequently the alleles conferring resistance were not completely concentrated in the most resistant cultivar studied (cv. Proctor). Heritability estimates obtained from F₂ and F₃ material suggested that field selection was of limited reliability, even when spreader drills were incorporated amongst the segregates. Single-plant selection (in F₂) was considered to be of little value, and the results of F_a head-row tests would require confirmation by replicated tests in subsequent generations.     

Race non-specific resistance to Bremia lactucae

Three ways of measuring race non-specific resistance to Bremia lactucae in lettuce were studied to determine the most important components of this form of resistance. Tests were conducted on leaf discs from 4–6 wk-old plants that were quantitatively inoculated and incubated under controlled conditions. Observations were made of disease intensity, latent period and production of conidia. Inoculum concentration markedly influenced disease intensity and latent period. Cv. Iceberg exhibited race non-specific resistance and this was characterised by a longer latent period, a lower disease intensity and reduced production of conidia compared to the susceptible control cv. Cobham Green. On older, intact plants, the differences in latent period and disease intensity between cultivars were even greater. Production of conidia was a particularly good criterion for discriminating levels of resistance.     

Components of partial resistance to Septoria nodorum in winter wheat

Components of partial resistance to Septoria nodorum were investigated in 10 cultivars of winter wheat having similar field resistances. The components measured were infection frequency, latent period, size, shape and rate of growth of lesions, spore production and its rate of increase. Latent period was found to be lognormally distributed. Some of the components of resistance were found to be significantly different between cultivars. Cluster analysis also showed that cultivars could be distinguished on the basis of their components of resistance. Principal components analysis indicated that resistance could be broken down into four underlying factors, three of which could be readily interpreted. The measurements of the components of resistance were combined in a model, the r-index, based on Van der Plank's r. The amount of variation between cultivars cast some doubt on the predictive value of the index but all the cultivar values were well within the range bounded by two ‘synthetic’ cultivars made up of combinations of either the most resistant or the most susceptible components. It is considered that the r-index has potential in screening for field resistance. The possibility of incorporating the most resistant-type components into one cultivar is discussed. The use of cultivar mixtures containing cultivars having similar field resistances is also discussed in the light of the variability found in this study.     

Components of Resistance to Early Blight in Four Potato Cultivars: Effect of Leaf Position

Components of early blight resistance were quantified in leaves of different ages in four potato cultivars. The components of resistance: incubation period (IP), lesion number (LN), early blight severity, lesion expansion rate (LER), latent period (LP) and spore production by lesion area (SPLA), were evaluated separately in the lower, middle and upper leaves of four potato cultivars. Plants of cultivar Aracy (resistant), Delta (moderately resistant), Desirée (susceptible) and Bintje (susceptible) were inoculated with an Alternaria solani isolate at the beginning of the flowering stage. Disease severity varied in different plant parts. In all cultivars, regardless of resistance, the smallest values of LN, and severity were recorded on the upper leaves, suggesting that young tissues are less susceptible. In cultivar Aracy, the IP was long, with small values of LN and LER and consequently, low values of early blight severity in all leaf positions were recorded. Although IP was long in cultivar Aracy, no differences between the moderately resistant cultivar Delta and the susceptible cultivars Bintje and Desirée could be detected for this component. The IP was only influenced by leaf position in cultivar Aracy. Clear differences in resistance levels among cultivars could be detected regarding LN, severity and LER. However, neither LP nor SPLA were associated with resistance level of cultivars or with leaf position. Analyses according to plant part suggest that evaluations on leaves of the middle third part are most suitable for screening for early blight resistance in potato.     

Inheritance of resistance to lettuce root aphid in the lettuce cultivars 'Avoncrisp' and 'Lakeland"

The inheritance of resistance in two lettuce cultivars to lettuce root aphid, Pemphigus bursarius, was studied in a series of laboratory and field experiments at Wellesbourne between 1989 and 1992. A source of total resistance in the cv. ‘Avoncrisp’ which is linked to the downy mildew resistance gene Dm6, was shown to be governed by a single dominant gene. There were no maternal effects evident in the inheritance of this resistance. The basis of the high level of resistance which exists in the cv. ‘Lakeland’ (formerly known as ‘Jubilee’) was also shown to be controlled by the same dominant gene. The linkage between Dm6 and root aphid resistance was broken in ‘Lakeland’ as this cultivar does not possess the Dm6 gene. The linkage was presumably broken when the original cross between the parents of cv. ‘Lakeland’, ‘Calmar’ and ‘Avoncrisp’ was made. Under laboratory conditions small numbers of aphids commence development on cv. ‘Lakeland’ but colonies fail to develop and under field conditions the resistance provides adequate Protectión against the pest. The resistance in both ‘Avoncrisp’ and ‘Lakeland’ was effective against a population of lettuce root aphid collected from an endive crop in southern France as well as being effective against the Wellesbourne population of this aphid.     

The phenotypic expression of QTLs for partial resistance to barley leaf rust during plant development

Partial resistance is generally considered to be a durable form of resistance. In barley, Rphq2, Rphq3 and Rphq4 have been identified as consistent quantitative trait loci (QTLs) for partial resistance to the barley leaf rust pathogen Puccinia hordei. These QTLs have been incorporated separately into the susceptible L94 and the partially resistant Vada barley genetic backgrounds to obtain two sets of near isogenic lines (NILs). Previous studies have shown that these QTLs are not effective at conferring disease resistance in all stages of plant development. In the present study, the two sets of QTL–NILs and the two recurrent parents, L94 and Vada, were evaluated for resistance to P. hordei isolate 1.2.1 simultaneously under greenhouse conditions from the first leaf to the flag leaf stage. Effect of the QTLs on resistance was measured by development rate of the pathogen, expressed as latency period (LP). The data show that Rphq2 prolongs LP at the seedling stage (the first and second leaf stages) but has almost no effect on disease resistance in adult plants. Rphq4 showed no effect on LP until the third leaf stage, whereas Rphq3 is consistently effective at prolonging LP from the first leaf to the flag leaf. The changes in the effectiveness of Rphq2 and Rphq4 happen at the barley tillering stage (the third to fourth leaf stages). These results indicate that multiple disease evaluations of a single plant by repeated inoculations of the fourth leaf to the flag leaf should be conducted to precisely estimate the effect of Rphq4. The present study confirms and describes in detail the plant development-dependent effectiveness of partial resistance genes and, consequently, will enable a more precise evaluation of partial resistance regulation during barley development.     

Components of quantitative resistance in sunflower to Alternaria helianthi

Components of quantitative resistance, spore production, incubation period, infection frequency and mean lesion size were measured in 17 sunflower accessions inoculated with conidia of Alternuria helianthi under controlled conditions. The same accessions were also rated for disease reaction in the field in 1994 and 1995 using a generated epidemic and varied in their disease reactions from highly susceptible to highly resistant. Spearman's ranking of accessions was highly correlated (r = 0.9) for both years; however, the ranking of components measured under controlled conditions with field severity was generally poor. Regression analysis of components with field severity ratings of the accessions showed that mean lesion size was highly correlated (r = O.74) and infection frequency was moderately correlated (r = 0.58) with the field severity ratings taken over the two years. Infection frequency was also well correlated (r = 0.75) with mean lesion size. Spore production and incubation period were poorly correlated with the field severity ratings for both years. An index based on infection frequency and mean lesion size gave a better correlation with the 1995 field severity ratings than either component alone, but in 1994 the index was not as well correlated with field severity as mean lesion size alone. It is suggested that mean lesion size, determined from plants 7–9 days after inoculation could be used to select for resistance to A. helianthi in the greenhouse. Infection frequency could also be used as a predictor of resistance, but to a lesser degree.     

Costs and benefits of hydroxamic acids-related resistance in winter wheat against the bird cherry-oat aphid, Rhopalosiphum padi L.

Estimations of infestation by the bird cherry-oat aphid (Rhopalosiphum padi) as well as measurements of grain yield in 26 Hungarian winter wheat cultivars under field conditions were correlated with the concentration of hydroxamic acids (Hx) in seedlings of those cultivars. The significant inverse relationship between infestation ratings and Hx levels in wheat showed that Hx, despite their decreased accumulation at later plant phenological stages, may be able to confer resistance against aphid infestation in the field. Since no significant relationship was found between grain yield and Hx levels in plants it is suggested that Hx accumulation does not impose a cost to the plant in terms of yield. These findings support earlier claims stressing the potential of Hx as breeding targets for aphid resistance in wheat.     

Compatibility of Erysiphe graminis Hybrids f. sp. secalis X f. sp. Tritici with Wheat Lines Involving Resistance Genes from Rye

Compatibility of hybrid cultures Erysiphe graminis ff. sp. secalis (SI) ×tritici (t2) was tested in the laboratory with wheat cultivars involving different resistance genes and with two rye cultivars. Segregation was observed on wheat without resistance gene and with resistance genes Pm1, Pm3b and Pm3c compatible with t2, but not on wheat with resistance gene Pm2, Pm 3a, Pm 4a and Pm 5 incompatible with t2, nor on rye. It was obvious that S1 involves avirulence genes to Pm1, Pm2, Pm 3a, pm 3b, Pm 3c, Pm 4a, Pm 5. Segregation was found on wheat cultivars involving rye resistance genes Pm 7 (Transfed) and Pm 8 (Kavkaz), but cv. Transec (Pm7) was incompatible with all cultures used, because Transec involves another gene for resistance. The results indicate that hybridization between formae speciales secalis and tritici of the fungus can be a source of fungus compatibility with wheat with rye resistance, even in field conditions.     

Further studies on the occurrence and inheritance of resistance in red raspberry to a resistance-breaking strain of raspberry bushy dwarf virus

A strain of raspberry bushy dwarf virus (RBDV-RB), discovered in England in 1981, readily infects by grafting many raspberry cvs that have gene Bu, which confers strong resistance or immunity to isolates of the common strain. Haida is one of two cultivars that are highly resistant or immune from RBDV-RB, but both its parents, cvs Creston and Malling Promise, are infectible. Studies of the segregation of resistance to both RBDV-RB and a common strain of RBDV (D200) in four progenies related to cv. Haida or its two parents, showed that resistance to RBDV-RB was heritable and occurred when gene Bu was present with a second resistance component whose inheritance is probably multigenic. There was some indication that the second component might be a form of partial resistance to graft inoculation of varying expression, and that cv. Haida possesses this resistance at a high level that has not been distinguished from immunity in the graft inoculations used. Cultivars Creston and Malling Promise possibly have this resistance to a lesser degree, while resistance in cv. Heritage has been distinguished from immunity only by extensive graft tests. Some possible implications for breeding RBDV-RB resistant cultivars are discussed.     

Partial resistance to rust (Puccinia allii) in cultivated leek (Allium ampeloprasum ssp.porrum): estimation and improvement

The extent to which resistance to leek rust (Puccinia allii) in commercially useful cultivars of leek (Allium ampeloprasum) could be developed by selection within current commercial cultivars or their recent predecessors was investigated. Differences in resistance among commercial cultivars was not great. However, a wide range of resistance levels was found in both half‐sib and inbred progeny derived from these cultivars. In a crop simulation, one of the breeding lines showed a reduction in the rate of new pustule appearance of 44% compared to cv. Wintra. It is concluded that selection within modern cultivar level material is a potentially productive strategy for the commercial development of leek rust resistance in the short to medium term.     

Inheritance of partial resistance to powdery mildew in spring wheat

Summary Four spring wheat (Triticum aestivum L.) cultivars exhibiting partial resistance to powdery mildew induced by Erysiphe graminis f.sp. tritici were crossed to a common susceptible cultivar to study the inheritance of resistance. The genetic parameters contributing to resistance were estimated by generation means analyses. Additive gene action was the most important genetic component of variation among generation means in all four crosses. Additive by additive effects were significant in one cross and both additive by additive and additive by dominance effects were significant in another. Dominance effects were not significant. The F2/F3 correlations in three crosses ranged from 0.27 to 0.43. Three additional crosses among resistant cultivars were employed to study the effectiveness of selection in improving resistance. By selecting the most resistant plants from the F2 and evaluating the progenies in the F4, increases in resistance ranging from 21% to 31% were obtained. In all crosses, there was transgressive segregation in both directions indicating that the genes conferring resistance to these cultivars differ and exhibit additive effects.     

Interactions between the diamondback moth, Plutella xylostella L. and glasshouse and outdoor-grown cabbage cultivars

Interactions between Brassica cultivars (cvs) and adult and larval stages of Plutella xylostella L. were examined. In six-way choice oviposition experiments, glasshouse-grown Chinese cabbage cv. Tip Top was preferred to five outdoor-grown temperate cabbage cultivars but was similar to four out of five tropical cultivars. Bionomic studies with neonate larvae on temperate cabbage cultivars showed that mean larval duration was longer and percentage survival less on older compared with younger plants but that plant age or cultivar had little effect on pupal weight. The fecundity of P. xylostella reared on outdoor-grown cabbage cultivars varied ten-fold between the poorest and the best cultivar but plant age had little effect on P. xylostella reared on Chinese cabbage. In a glasshouse experiment, survival of P. xylostella on Chinese cabbage was significantly greater compared with field-grown glossy-leaved, normal bloom green cabbage and red cabbage. In bioassays with neonate larvae, when the leaves were placed vertically instead of horizontally a significant level of intrinsic resistance was revealed for glasshouse and field-grown glossy-leaved cabbage but not for Chinese cabbage. Observed differences in host status to P. xylostella are discussed in relation to the potential for partial plant resistance in control programmes.