Incomplete specific resistance to Bremia lactucae in lettuce

The complementary pair of dominant genes. Dm 7/1 and Dm 7/2 and the single dominant gene Dm 6 for specific resistance in Bremia lactucae in lettuce, condition incomplete resistance to non-virulent (incompatible) isolates of B. lactucae in both seedling and mature tissue. This incomplete resistance is characterised by a marked reduction in spotophore production compared with fully susceptible (compatible) race/cultivar combinations and by macroscopically visible hypersenstive cell necrosis. The incomplete resistance. In seedling the resistance conditioned by genes giving more complete resistance. In seedlings the resistance conditioned by genes Dm 7/1 and 7/2 was incomplete in outdoor 'crisp' lettuce genotypes and the resistance conditioned by gene Dm 6 was only completely expressed at 20°C.     

The influence of seedling age and development on the infection of lettuce by Bremia lactucae

Laboratory methods are described for the culture and observation of Bremia lactucae on lettuce. Mycelial growth of the fungus within the cotyledons of the cultivar Cheshunt Early Giant progressed to a minimum density at which sporulation occurred, and then continued to a maximum density coinciding with peak sporulation. The timing and intensity of sporulation was influenced by the concentration of the inoculum. Seedlings often became systemically infected and this, together with intensity of sporulation, was a good indicator of fungal growth and hence host susceptibility. The severity of disease development on four susceptible lettuce cultivars (Cheshunt Early Giant, Paris White, Trocadero Improved and Crackling Ice) decreased with age and development of the seedlings at inoculation. The reasons for this are discussed.     

Linkage between genes for resistance to downy mildew (Bremia lactucae) in lettuce

Data are presented for the joint segregation of the gene combinations: Dm-2/Dm-3, Dm-3/Dm-6, Dm-2/Dm-6 and Dm-6/Dm-8, following inoculation with five races of B. lactucae. It is postulated that Dm-2, 3 and 6 comprise a tight linkage group but that Dm-6 and Dm-8 are not linked as has been proposed previously. Information on the reaction of some resistant lettuce cultivars to certain B. lactucae races and some new postulated host genotypes is also presented.     

Studies on a method of carry-over of downy mildew (Bremia lactucae) of lettuce

Summary Bremia lactucae produced oospores in the decayed stem tissue of infected lettuce plants. These oospores caused cotyledon and first leaf infections ofBremia lactucae when added to the rooting medium of lettuce seedlings grown under sterile conditions.     

The hypersensitive reaction, membrane damage and accumulation of autofluorescent phenolics in lettuce cells challenged by Bremia lactucae

The expression of resistance to Bremia lactucae determined by the resistance genes Dm5/8 and Dm7 in lettuce was examined; incompatibility involved the hypersensitive reaction (HR) which occurred only within penetrated cells at early and late stages of fungal development, respectively. Autofluorescence observed under UV and blue light excitation in cells undergoing the HR was associated with the accumulation of ester-linked syringaldehyde and caffeic acid on plant cell walls. Two phases of phenolic deposition were identified. The first was highly localized around penetration points and occurred during incompatible and compatible interactions. The second and major phase was only activated after the occurrence of irreversible membrane damage in the penetrated cell and was reduced by inhibitors of mRNA synthesis. Fungal structures, primary and secondary vesicles, intercellular hyphae and haustoria also became autofluorescent during incompatible interactions. Changes in the fluorescence due to preformed phenolics located in the plant cell vacuole were found just before plasma membrane damage became irreversible during the HR. In addition to localized deposition of phenolics, increases in the concentrations of the major free phenolic esters identified as dicaffeoyl tartaric and chlorogenic acids also occurred during incompatible interactions. The results suggest that membrane damage in penetrated cells occurs at different rates in resistance controlled by Dm5/8 and Dm7 and indicate an important role for irreversible membrane damage in lettuce as a key signalling event leading to widespread activation of defence responses in surrounding cells.     

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.     

Race specific resistance to lettuce downy mildew (Bremia lactucae) in the lettuce cultivar Bourguignonne Grosse Blonde d'Hiver

A stock of lettuce cv. Bourguignonne Grosse Blonde d'Hiver used in previous studies was shown to carry the race specific resistance factor R1 and not R9 as originally thought. Investigations into a commercial stock of cv. Bourguignonne which exhibited a heterogeneous reaction to some B. lactucae isolates showed that it was a mixture of two lines. Each line carried one of two R-factors (referred to as R9A and R9B) in a homozygous condition, although their identification was made difficult by the presence of R5/8. No plants were found which carried both R9A and R9B.     

Virulence of Bremia lactucae in Sweden and race-specific resistance of lettuce cultivars to Swedish isolates of the fungus

Virulence surveys of Swedish Bremia lactucae populations confirmed that the virulence factors vl to v12 were present in high or very high frequencies. Virulence associated with recently defined new resistance genes was also present. Laboratory tests of lettuce cultivars and Lactuca accessions using different Bremia isolates and field tests with natural inoculum showed that previously undetected virulence factors were present. Due to a lack of highly effective genes for specific resistance and the frequent sexual recombination of virulence genes it is suggested that any future breeding programmes concentrate on non-specific resistance.     

The genetics of race specific resistance in lettuce (Lactuca sativa) to downy mildew (Bremia lactucae)

Data are presented on the segregation of resistance to five British races and two Dutch races of Bremia lactucae in the F₂ progenies of crosses involving seven resistant and several susceptible lettuce cultivars. These data and also those previously published by other workers are considered in relation to the systematic model proposed by Crute & Johnson (1976) to explain the genetics of race specific resistance to B. lactucae in lettuce. It is shown that, with minor modifications, the model accommodates almost all of the previously published data and correctly predicts the new data, except for one set which cannot at present be interpreted. It is concluded that genetic evidence exists for the presence, among various cultivars of lettuce, of at least four and possibly five different dominant resistance genes of major effect designated Dm₂, Dm₃, Dm₄, Dm₆ and Dm₈; and of a pair of dominant genes with complementary effect designated Dm7/1 and Dm7/2. The resistance conferred by these genes is specified in relation to five British races, five Dutch, three Israeli and one United States race of the fungus. Resistance genotypes are proposed for cultivars Avoncrisp, Avondefiance, Calmar, Great Lakes 659, Kares, Meikoningen, Mildura, Proeftuins Blackpool, Solito, Valverde, Ventura and the USDA line PI 164937.     

Laboratory and field tests of zineb and other fungicides against downy mildew of lettuce (Bremia lactucae Regel)

In laboratory tests on young lettuce plants, the protection provided by sprays of zineb wettable powder against artificial inoculations with Bremia lactucae was less than that obtained with tank-mix zineb, and with several other dithiocarbamates and related fungicides. Experiments on autumn lettuce in the field also showed that tank-mix zineb was more effective than the wettable powder. On winter lettuce in frames, however, zineb wettable powder gave control similar to or better than that given by any of the other materials tested. Infection in most of these outdoor experiments was not heavy. None of the fungicides gave very high control or markedly increased the number of hearted or marketable plants. When zineb wettable powder was sprayed on autumn lettuce at the rate of 2 lb./acre it gave similar disease reductions whether applied in low, medium or high volumes of water.     

Linkage analysis of genes for resistance to downy mildew (Bremia lactucae) in lettuce (Lactuca sativa)

Summary The genetics of specific resistance was studied in F₂ populations which segregated for either one or two resistance genes. The resistance factors 1, 11 and 14 which had not previously been characterized genetically segregated as single dominant genes (Dm). Resistance was determined by three linkage groups; R 1/14, 2, 3, and 6 in the first, R 5/8, and 10 in the second and R 4, 7 and 11 in the third. Cultivars of lettuce commonly used in the differential series to detect virulence to R3 and R10, were demonstrated to carry two tightly linked resistance genes. Implications of this linkage arrangement to the manipulation and characterization of these resistance genes are discussed.     

A genetic map of the lettuce downy mildew pathogen,Bremia lactucae,constructed from molecular markers and avirulence genes

The genetic map of Bremia lactucae was expanded utilizing 97 F(1) progeny derived from a cross between Finnish and Californian isolates (SF5xC82P24). Genetic maps were constructed for each parent utilizing 7 avirulence genes, 83 RFLP markers, and 347 AFLP markers, and a consensus map was constructed from the complete data set. The framework map for SF5 contained 24 linkage groups distributed over 835cM; the map for C82P24 contained 21 linkage groups distributed over 606cM. The consensus map contained 12 linkage groups with markers from both parents and 24 parent-specific groups. Six avirulence genes mapped to different linkage groups; four were located at the ends of linkage groups. The closest linkages between molecular markers and avirulence genes were 3cM to Avr4 and 1cM to Avr7. Mating type seemed to be determined by a single locus, where the heterozygote determined the B(2) type and the homozygous recessive genotype determined the B(1) type.     

Occurrence of Bremia lactucae in Natural Populations of Lactuca serriola

In the period 1996–2001 the natural occurrence of Bremia lactucae (lettuce downy mildew) on Asteraceae plants was studied in the Czech Republic. Lactuca serriola (prickly lettuce) is the most common naturally growing host species of B. lactucae. Infection of plants was recorded during the whole vegetation season with the first occurrence in April and last in November. Bremia lactucae was found on host plants in all developmental stages. High percentages of naturally infected populations of L. serriola were recorded. Host plants exhibited broad variation in phenotypic expression of disease symptoms and degree of infection, however, the intensity of infection was rather low in the majority of populations. Geographic distribution of B. lactucae was studied in the two main parts of Czech Republic, central and southern Moravia, and eastern, northern and central Bohemia. Bremia lactucae was recorded in all these areas. Nevertheless, in the warmest parts of the Czech Republic (southern Moravia) only sporadic occurrence of the pathogen was recorded. Bremia lactucae infection on L. serriola and disease severity was judged also in relation to the type of habitat, and the size and density of host plant populations. However, no substantial differences among various habitats were found; only host plants growing in urban areas were frequently free of infection and the degree of infection was very low. Nevertheless, these plants were commonly infected with powdery mildew (Golovinomyces cichoracearum), which is most aggressive pathogen of this type of habitat.