Partial resistance in lettuce to downy mildew (Bremia lactucae). 1. Search for partially resistant genotypes and the influence of certain plant characters and environments on the resistance level

Summary In three bicyclic inoculation experiments in the field 756 genotypes from the IVT Lactuca gene bank were screened for differences in partial resistance against Bremia lactucae. Significant differences between genotypes were found. No linkage was found between the presence or absence of R-genes and partial resistance nor was there a clear relationship between plant characters such as size, erectness and lettuce type and resistance level. Comparison of test results for genotypes which were tested twice and in different years and/or environments revealed a rather good relationship. So no clear genotype × environment interactions for resistance level were found. Results imply that the bicyclic inoculation experiments are useful to determine levels of partial resistance.     

The identification and characteristics of field resistance to lettuce downy mildew (Bremia lactucae Regel)

Summary Eighty-one accessions of three Lactuca species which showed no recognisable race specific resistance to Bremia lactucae when tested in the laboratory as seedlings, exhibited different degrees of susceptibility when exposed to natural field infection. As a group, crisp genotypes had less mildew and a slower rate of disease development than other types of lettuce. Wild forms of Lactuca sativa and Lactuca serriola were particularly susceptible. In a further trial, the low field susceptibility of three lettuce cultivars (Iceberg, Batavia blonde de Paris and Grand Rapids) was confirmed. Disease development on cv. Iceberg was compared to that on the highly susceptible cv. Hilde in experiments where the two cultivars were grown either in close proximity or in isolation. The absolute level of attack on cv. Iceberg depended upon disease pressure and differences between the two types only became apparent approximately 8 wk after sowing.     

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.     

New sources of major gene resistance in Lactuca to Bremia lactucae

Summary A total of 1789 accessions of several lettuce collections was screened to find new major gene resistance to the downy mildew fungus Bremia lactucae Regel. The accessions belonged to the species Lactuca sativa (N=1288), L. serriola (N=399), L. saligna (N=52) and L. virosa (N=50). A total of 20 races of B. lactucae were used, 14 of which were NL-races, isolated from cultivated lettuce in the Netherlands. The other six races were isolated from wild L. serriola in Czechoslovakia. The accessions were initially screened with two races: NL1 and NL3. Accessions with resistance to one or both of these races were tested with the other races. Phenotypes with new resistance were found in accessions of all four Lactuca species. Of L. sativa, four accessions were found with resistance phenotypes that could not be explained by combinations of known major genes. Many accessions of L. serriola had resistance phenotypes that indicated the presence of unknown resistance genes. All interactions between accessions of L. saligna and races of B. lactucae were incompatible in leaf disc tests, except for four accessions, which showed some sporulation with race NL6. Several accessions of L. virosa were resistant to all races used. Other accessions of L. virosa gave a race-specific interaction with B. lactucae.     

Inheritance of resistance to Zucchini yellow mosaic virus in the interspecific cross Cucurbita maxima × C. ecuadorensis

Summary More than 200 accessions of three wildLactuca species were screened in the laboratory for race-specific resistance toBremia lactucae. Only a fewLactuca entries showed resistance both as seedlings and in a leaf disc test. Accessions ofL. serriola andL. sativa had a low degree of resistance. The best entries were also compared under field conditions. TheL. saligna entries were totally free from disease in the field test. High resistance was also recorded in a fewL. serriola entries and in lettuce cultivars such as Saffier and Mariska.     

The location and characteristics of novel sources of resistance to Bremia Lactucae Regel (downy mildew) in wild Lactuca L. Species

Summary Twenty-one accessions of 3 wild Lactuca species which could be hybridised with L. sativa, the cultivated lettuce, were inoculated at different stages of plant development with 3 multivirulent isolates of Bremia lactucae. Nineteen sources of resistance to B. lactucae, not attributable to the previously recognised resistance factors 1–11 were identified. Two lines of L. serriola showed similar resistance patterns as lines carrying R11. The resistance of some accession was incomplete particularly at the seedling stage and this phenomenon may be race specific.Tests on segregating F₂ populations of crosses between 2 different L. serriola accessions and L. sativa cultivars showed that the resistance in one line (LSE/18) appears to be inherited as a single dominant gene, which is sometimes incomplete in expression and allelic to either Dm6 or R7. The segregation patterns for resistance in PI 281876 did not give readily interpretable ratios.To assess the frequency of occurrence in B. lactucae populations of virulence factors to overcome this novel resistance, 11 of the novel sources of resistance were inoculated with numerous collections of the pathogen from the UK, Czechoslovakia and elsewhere and found to show a high level of resistance.     

An association between resistance to root aphid (Pemphigus bursarius L.) and downy mildew (Bremia lactucae Regel) in lettuce

Summary Many lettuce cultivars (Lactuca sativa L.) with high resistance to lettuce root aphid (Pemphigus bursarius L.) also carried the gene Dm-6 for specific resistance to downy mildew (Bremia lactucae Regel). This suggests the possibility of linkage between this gene and root aphid resistance. The origin of this association is discussed.     

Effects of inoculum density and temperature on three components of leaf rust resistance controlled by Lr34 in wheat

Summary Different inoculum densities had negligible effects on latent period, uredinium density and uredinium size measured on flag leaves of adult RL6058 (Thatcher^*6/PI58548[Lr34]) plants kept at low (17.5°C) post-infection temperatures in a glasshouse. In a qualitative assessment of rust severity at higher (24.6°C) temperatures, all three components of resistance indicated a susceptible flag leaf response on RL6058. In the latter environment, precise estimations of receptivity to different inoculum densities showed that adult RL6058 plants supported significantly less pustules than the leaf rust-susceptible cultivar Thatcher. In tillering plants, statistically equal numbers of uredinia developed on RL6058 and Thatcher in all paired temperature-inoculum density combinations. Growth stage-related susceptibility, and higher temperatures conducive to a shorter latent period and larger uredinia, could result in high terminal severities of Puccinia recondita f. sp. tritici on wheat genotypes containing Lr34. The reduction in receptivity associated with this gene may contribute, however, to delayed disease increase on cultivars or lines with monogenic Lr34 resistance.     

The inheritance and biochemistry of resistance to Sclerotinia sclerotiorum leaf infections in sunflower (Helianthus annuus L.)

Summary Resistance of sunflower leaves to attack by Sclerotinia sclerotiorum was studied by infecting them, in the field, with agar disks containing Sclerotinia mycelium. Resistance levels were determined by the length of lesions after a given period. There were significant differences in reaction between both sunflower hybrids and inbred lines. Different Sclerotinia isolates gave the same classification of sunflower genotypes. The results of trials repeated in one year or different years were significantly correlated. The general combining ability variance/specific combining ability variance ratio was 1.35. Strict sense heritability was 0.61±0.03. The midparent-offspring correlation coefficient was significant, but the relation between per se values of the male parents used with the values of their hybrids was much closer than that for the female parents. Mean heterosis for resistance was 28.9%, compared with the midparent. The results of the leaf resistance test are frequently correlated with levels of resistance to root attack by Sclerotinia. Possible use of this test in breeding both directly for leaf resistance and indirectly for root resistance are discussed. H.P.L. Chromatography studies of the phenols present in healthy and infected leaves distinguished 19 compounds, all of the inhibitin type. There was a large increase in phenol content in leaves infected by Sclerotinia, for all genotypes. However, more especially in uninfected leaves, the contents of 3 chromatogram peaks, numbered 4, 6 and 9, showed a close relation with levels of Sclerotinia resistance. It is proposed that these compounds could be used as markers of certain types of resistance.     

Partial resistance in lettuce to downy mildew (Bremia lactucae). 3. Correspondence between resistance levels of cotyledons and leaf discs and resistance of adult plants

Summary The correlation between resistance levels of seedlings with cotyledons only and leaf discs taken from young and old leaves on the one hand and the resistance of adult plants in bicyclic field tests on the other was investigated. Components of resistance were: latent period (LP) the percentage of seedlings (SEED) or the number of cotyledons (COT) infected, the number of sporangiophores (SPOR) per cotyledon or leaf dise and the % of the surface (SURF) per cotyledon or disc covered with sporangiophores.For both cotyledons and leaf discs, significant differences occurred between plant genotypes for LP, SPOR and SURF. The correspondence between results of cotyledon testing and field testing was generally negligible. There was a rather good correlation between the resistance levels of leaf discs, expressed in SPOR and SURF and the resistance of adult plants. On the basis of a smaller coefficient of variation for SPOR this parameter should be preferred to predict the resistance of adult plants by testing leaf discs.     

Partial resistance of barley to leaf rust,Puccinia hordei. V. Analysis of the components of partial resistance in eight barley cultivars

Summary Eight spring barley cultivars, respresenting the known range in partial or slow rusting resistance to leaf rust, Puccinia hordei, were investigated for their effects on the components of partial resistance; infection frequency, latent period, infectious period and spore production per uredosorus per day. Considerable variation was observed among the cultivars for each of the components. The cultivar effects on the components tend to be associated. Cultivar L94 for instance, shows the highest infection frequency, the shortest latent period and a long infectious period. Julia and Vada both have a low infection frequency, a long latent period and a low spore production per sorus per day. This association, though, is only a partial one.The total spore production per unit leaf area (the combined result of the four components) appeared highly correlated with the partial resistance in the field (r=0.85). Only a relatively small portion of the variation in partial resistance cannot be explained by the four components studied. Several other aspects, which might affect the rate of epidemic development, are discussed.Latent period, measuring the onset of the new spore production, estimated partial resistance as well as total spore production did (r=–0.85). In order to evaluate the partial resistance of barley genotypes in the greenhouse the latent period is preferred above total spore production as it is measured more easily, more accurately and sooner after inoculation.     

The role of resistance breeding in the intergrated control of downy mildew (Bremia lactucae) in protected lettuce

Summary Over the last 30 years, six resistance alleles (Dm2, Dm3, Dm6, Dm7, Dm11 andDm16) located in two linkage groups, have contributed to the control of downy mildew in lettuce crops grown under protection (glass or polythene) in northern Europe. More recently, an as yet genetically uncharacterised resistance factor, R18, has also begun to assume importance. The occurrence of the various combinations of these resistance alleles that exist in commercial cultivars has been dictated by the pathotypes ofBremia lactucae used in their selection but also restricted by linkage in repulsion. In the UK, a pathotype ofB. lactucae insensitive to phenylamide fungicides, such as metalaxyl, emerged in 1978 and became prevalent throughout lettuce production areas in subsequent years. The specific virulence of this pathotype was identical to the previously described phenylamide sensitive pathotype NL10 and cultivars carryingDm11, Dm16 or R18 were resistant. Consequently, an integrated control strategy based on the utilisation of metalaxyl on cultivars carryingDm11 provided effective control in UK until 1987 when a new phenylamide insensitive pathotype began to cause problems. The specific virulence of this second pathotype, which was first reported in the Netherlands and France, was identical to the previously described phenylamide sensitive pathotype NL15. Cultivars carryingDm6, Dm16 or R18, but notDm11, were resistant to NL15; consequently an appropriate change in the cultivar recommendations for use in the integrated control strategy was successfully promulgated. It is predicted that variations of this integrated control strategy involving the use of appropriately selectedDm gene combinations may prove effective for some time. This prediction is based on studies of the status of the avirulence loci in the two phenylamide insensitive pathotypes and of the specific virulence characteristics of phenylamide sensitive components of the pathogen population.     

Diallel analysis of barley for resistance to leaf stripe and impact of the disease on genetic variability for yield components

Barley breeders in Syria attempting to develop barley (Hordeum vulgare L.) cultivars resistant to barley leaf stripe (BLS) disease caused by Pyrenophora graminea Ito & Kuribayashi [anamorph Drechslera graminea (Rabenh. Ex. Schlech. Shoem.)]. Information on the combining ability for BLS resistance in Syria is not available. This study was conducted to evaluate, in 10 genetically diverse barley parents, general combining ability (GCA) and specific combining ability (SCA) effects towards the determination of the genetic basis of disease resistance and to estimate genetic variability for yield components and its modification by BLS. Ten parental genotypes varying in their reactions to BLS were crossed in a half-diallel mating design to generate 45 full-sib families. The families and the parents were inoculated with P. graminea and evaluated for resistance in replicated field tests (three inoculated and three non-inoculated plots). The parents chosen showed wide variations for resistance to BLS. Genetic component analysis showed significant effects for both GCA and SCA for resistance to BLS, suggesting that additive as well as non-additive genetic mechanisms were involved in the expression of resistance in these parents. GCA effects were more important than SCA effects. Resistant parents exhibited high negative GCA indicating that additive gene effects were more predominant, and suggesting their prime suitability for use in barley breeding programs to improve resistance to BLS. Narrow-sense heritability was 58% and broad-sense heritability was 99% indicating that selection for BLS resistance should be effective in these crosses. A high genetic variability for the agronomic traits studied was observed. Yield components decreased significantly in inoculated plants and more pronounced in diseased plants. Significant GCA was observed for all traits. Values for GCA were, in some cases, significantly modified by BLS. This indicates that attention must be paid to the danger of drawing conclusion in quantitative genetics studies dealing with both diseased and healthy plants. Two genotypes, Banteng and Igri, had high negative GCA effects and are promising parents for enhancement of BLS resistance.     

Identification of growth stage dependent expression of partial resistance of barley to powdery mildew

Summary Genotypic differences for growth stage dependent expression of partial resistance to barley powdery mildew have been identified on the basis of two components of resistance in the glasshouse and the pattern of epidemic development in the field.Differences for infection frequency and proportion of sporulating colonies were highly significant between the 16 genotypes investigated at four stages of plant development in the glasshouse. Both resistance components were significantly correlated mutually (r=0.73 to 0.86) and with the infection level on leaves developed at comparable growth stages in the field (r=0.52 to 0.73). The infection level of seedlings in the glasshouse was significantly correlated with the infection level of the leaves of the first until the third node in the field (r=0.70 to 0.73). Adult plant resistance was predominantly expressed at the uppermost leaf. A clear genotype × growth stage interaction was apparent for infection level; some genotypes showed partial resistance predominantly at the seedling stage and others predominantly at the adult plant stage.Abbreviations IF Infection Frequency, number of colonies per cm² leaf area, glasshouse experiment - IL Infection Level, number of colonies per cm² leaf area, field experiment - PSC Proportion Sporulating Colonies from total number of colonies, glasshouse experiment     

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.     

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.     

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.     

Evaluation of globe artichoke and related germplasm for resistance to artichoke latent virus

Summary A total of 280 globe artichoke (Cynara scolymus L.) clones were assessed for reaction to naturally occurring artichoke latent virus (ALV) in field trials. Also, the reaction of 17 seed progenies of crosses which produced ALV-resistant clones in the field, 20 self progenies of parental genotypes common to these crosses, self progenies of 17 artichoke cultivars, and 7 accessions ofC. cardunculus andC. syriaca were evaluated in the glasshouse following mechanical inoculation with ALV. The local and systemic presence of ALV, and the relative virus content in infected plants were determined by ELISA.All cultivars and most of the otherC. scolymus material tested (clones and seed progenies) proved to be highly susceptible to ALV. Ten progenies were less susceptible to mechanical inoculation as shown by the moment the virus could be detected, and a slow rate of accumulation of viral antigen. Some plants in two seed progenies possessed apparently a high resistance as could be concluded from the failure of ALV to spread or to multiply in plant.Accessions ofC. cardunculus andC. syriaca were also highly susceptible to ALV. However, an interspecific hybridC. cardunculus x C. syriaca presented the highest level of resistance, the plants remained free of local and systemic infection. Ratios of resistant to susceptible plants in self progenies of resistant plants suggested that this resistance was dominant.     

Genetic basis of pod borer (Helicoverpa armigera) resistance and grain yield in desi and Kabuli Chickpea (Cicer arietinum L.) under unprotected conditions

A series of half-diallel crosses involving early, medium and late maturity desi and kabuli type chickpea (Cicer arietinum L.) genotypes with stable resistance to Helicoverpa pod borer, along with the parents, were evaluated at two locations in India to understand the inheritance of pod borer resistance and grain yield. Inheritance of resistance to pod borer and grain yield was different in desi and kabuli types. In desi type chickpea, the additive component of genetic variance was important in early maturity and dominance component was predominant in medium maturity group, while in the late maturity group, additive as well as dominance components were equally important in the inheritance of pod borer resistance. Both dominant and recessive genes conferring pod borer resistance seemed equally frequent in the desi type parental lines of medium maturity group. However, dominant genes were in overall excess in the parents of early and late maturity groups. In the kabuli medium maturity group, parents appeared to be genetically similar, possibly due to dispersion of genes conferring pod borer resistance and susceptibility, while their F₁s were significantly different for pod borer damage. The association of genes conferring pod borer resistance and susceptibility in the parents could be attributed to the similarity of parents as well as their F₁s for pod borer damage in kabuli early and late maturity groups. Grain yield was predominantly under the control of dominant gene action irrespective of the maturity groups in desi chickpea. In all the maturity groups, dominant and recessive genes were in equal frequency among the desi parental lines. Dominant genes, which tend to increase or decrease grain yield are more or less present in equal frequency in parents of the early maturity group, while in medium and late maturity groups, they were comparatively in unequal frequency in desi type. Unlike in desi chickpea, differential patterns of genetic components were observed in kabuli chickpea. While the dominant genetic component was important in early and late maturity group, additive gene action was involved in the inheritance of grain yield in medium duration group in kabuli chickpea. The dominant and recessive genes controlling grain yield are asymmetrically distributed in early and medium maturity groups in kabuli chickpea. The implications of the inheritance pattern of pod borer resistance and grain yield are discussed in the context of strategies to enhance pod borer resistance and grain yield in desi and kabuli chickpea cultivars.     

Alternative methods of interpreting measurements of resistance to the leafhopper Empoasca kraemeri Ross and Moore in the common bean,Phaseolus vulgaris L.

Summary Appropriate definitions of, and methods of measuring, resistance are sought using Phaseolus vulgaris, a major source of protein in Latin America, and its most important pest, Empoasca kraemeri. The concepts of general vigour and narrow-sense resistance are defined: these two components make up broad-sense resistance. The validity of these definitions, which depend on the inclusion of tolerance as a resistance mechanism, is discussed. In order to distinguish the components, 38 genotypes of P. vulgaris were grown with and without insecticidal proctection, in three replications in a split-plot design, with three planting dates. Seed yield per plant, and a visual score of damage symptoms or vigour, were measured. Three methods of estimating narrow sense resistance are considered: 1) the genotype × insecticide interaction effect in a split-plots analysis, 2) the yield of each genotype unprotected (Y_u) divided by its yield protected (Y_p). and 3) the deviation of each genotype from a regression of Y_u on Y_p. The third method overcomes some objections to the first two, and can be applied to damage and vigour scores as well as to yields. The 38 genotypes differed in narrow-sense resistance according to all three methods, and it appears that in practice Methods 2 and 3 were both valid. They also differed in general vigour. Breeders working with many pests and crops have found that low damage and high yield in the presence of the pest are effective selection criteria. The work reported here supports this view.