Structure and variation in the wild-plant pathosystem: Lactuca serriola–Bremia lactucae

Abstract

Over the past decade, extensive research on the wild-plant pathosystem, Lactuca serriola (prickly lettuce)–Bremia lactucae (lettuce downy mildew), has been conducted in the Czech Republic. Studies focused on pathogen occurrence and distribution, host range, variation in symptom expression and disease severity, interactions of B. lactucae with another fungal species (Golovinomyces cichoracearum) on L. serriola, variation in resistance within natural populations of L. serriola, the structure and dynamics of virulence within populations of B. lactucae, sexual reproduction of B. lactucae, and a comparison of virulence structure and changes in B. lactucae populations occurring in wild (L. serriola) and crop (L. sativa) pathosystems. The incidence of B. lactucae on naturally growing L. serriola and other Asteraceae was recorded. Lactuca serriola was the most commonly occurring host species, followed by Sonchus oleraceus. Over the duration of these studies, the incidence of B. lactucae in L. serriola populations varied between 45–87%. Disease incidence and disease prevalence were partly related to the size, density and different habitats of L. serriola populations. In addition to B. lactucae infection, infection by the lettuce powdery mildew fungus (Golovinomyces cichoracearum) was quite common, including co-infection. Variation in resistance to B. lactucae was studied by using ten isolates (NL and BL races with known virulence patterns) at a metapopulation level, i.e. 250 L. serriola samples representing 16 populations from the Czech Republic (CZ). Our comparisons revealed broad variation in host resistance among host populations and also intrapopulation variability. In the CZ populations, 45 resistance phenotypes were recorded, the most frequent were race-specific reaction patterns. Structural and temporal changes in virulence variation of B. lactucae populations on L. serriola were studied during 1998–2005. Altogether, 313 isolates of B. lactucae originating from the Czech Republic were examined for the presence of 32 virulence factors (v-factors), and 93 different virulence phenotypes (v-phenotypes) were recorded. A study of v-factor frequency showed that common v-factors in B. lactucae populations match some of the race-specific resistance genes/factors (Dm genes or R-factors) originating from L. serriola. The highest frequency was recorded by v-factors v7, v11, v15–17, and v24–30. In contrast, v-factors (e.g. v1–4, 6, and 10) matching Dm genes originating from L. sativa were very rare. This demonstrates the close adaptation of B. lactucae virulence to the host (L. serriola) genetic background. Temporal changes in virulence frequencies over the period were recorded. In many v-factors (v11, v14, v16, and v25–28), fluctuations were observed, some (v14 and v17) shifting to higher frequencies, and others (v5/8 and v23) decreasing. The occurrence of mating types was studied (1997–1999) in a set of 59 B. lactucae isolates. Both compatibility types (B1 and B2) were recorded; however the majority of the isolates (96%) were type B2. A comparative study of B. lactucae virulence variation between the wild (L. serriola) and crop (L. sativa) pathosystems showed major differences. Migration and gene flow between both pathosystems and the potential danger of wild B. lactucae populations for cultivated lettuce are discussed. This paper summarizes comprehensive and unique research on an oomycete pathogen (B. lactucae) that is shared between a crop (lettuce, L. sativa) and its close wild relative (prickly lettuce, L. serriola). The data demonstrate clear evidence about race-specific interactions, variation and changes in virulence, and coevolutionary relationships in the wild pathosystem L. serriola–B. lactucae. Conclusions contribute to the broadening and better understanding of gene-for-gene systems in natural host–pathogen populations and their relationships to crop pathosystems.