Systemic acquired resistance is a pathogen-inducible defense mechanism in plants. The resistant state is dependent on endogenous accumulation of salicylic acid (SA) and is characterized by the activation of genes encoding pathogenesisrelated (PR) proteins. Recently, selected nonpathogenic, root-colonizing biocontrol bacteria have been shown to trigger a systemic resistance response as well. To study the molecular basis underlying this type of systemic resistance, we developed an Arabidopsis-based model system using Fusarium oxysporum f sp raphani and Pseudomonas syringae pv tomato as challenging pathogens. Colonization of the rhizosphere by the biological control strain WCS417r of R fluorescens resulted in a plant-mediated resistance response that significantly reduced symptoms elicited by both challenging pathogens; Moreover, growth of R syringae in infected leaves was strongly inhibited in R fluorescens WCS417r-treated plants. Transgenic Arabidopsis NahG plants, unable to accumulate SA, and wild-type plants were equally responsive to R fluorescens WCS417r-mediated induction of resistance. Furthermore, R fluorescens WCS417r-mediated systemic resistance did not coincide with the accumulation of PR mRNAs before challenge inoculation. These results indicate that R fluorescens WCS417r induces a pathway different from the one that controls classic systemic acquired resistance and that this pathway leads to a form of systemic resistance independent of SA accumulation and PR gene expression.

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It is frequently hypothesised that high soil fungal/bacterial ratios are indicative for more sustainable agricultural systems. Increased F =B ratios have been reported in extensively managed grasslands. To determine the shifts in fungal/bacterial biomass ratio as influenced by grassland management and to find relations with nitrogen leaching potential, we sampled a two-year-old field experiment at an organic experimental farm in the eastern part of The Netherlands. The effect of crop (grass and grass-clover), N application rate (0, 40, 80, 120 kg N ha À1 ) and manure type (no manure, farm yard manure and slurry) on the F=B ratio within three growing seasons was tested, as well as relations with soil and crop characteristics, nitrate leaching and partial N balance. Biomass of fungi and bacteria was calculated after direct counts using epifluorescence microscopy. Fungal and bacterial biomass and the F =B ratio were higher in grass than in grassclover. The F =B ratio decreased with increasing N application rate and multiple regression analysis revealed a negative relationship with pH. Bacterial activity (measured as incorporation of [ H]thymidine and [14 C]leucine into bacterial DNA and proteins) showed the exact opposite: an increase with N application rate and pH. Leaching increased with N application rate and was higher in grass-clover than in grass. Partial N balance was more positive at a higher N application rate and showed an inverse relationship with fungal biomass and F =B ratio. We conclude that the fungal/bacterial biomass ratio quickly responded to changes in management. Grasslands with higher N input showed lower F =B ratios. Grass-clover had a smaller fungal biomass and higher N leaching than grass. In general, a higher fungal biomass indicated a lower nitrogen leaching and a more negative partial N balance (or smaller N surplus), but more observations are needed to confirm the relationship between F =B ratio and sustainability. r

Systemic acquired resistance is a pathogen-inducible defense mechanism in plants. The resistant state is dependent on endogenous accumulation of salicylic acid (SA) and is characterized by the activation of genes encoding pathogenesisrelated (PR) proteins. Recently, selected nonpathogenic, root-colonizing biocontrol bacteria have been shown to trigger a systemic resistance response as well. To study the molecular basis underlying this type of systemic resistance, we developed an Arabidopsis-based model system using Fusarium oxysporum f sp raphani and Pseudomonas syringae pv tomato as challenging pathogens. Colonization of the rhizosphere by the biological control strain WCS417r of R fluorescens resulted in a plant-mediated resistance response that significantly reduced symptoms elicited by both challenging pathogens; Moreover, growth of R syringae in infected leaves was strongly inhibited in R fluorescens WCS417r-treated plants. Transgenic Arabidopsis NahG plants, unable to accumulate SA, and wild-type plants were equally responsive to R fluorescens WCS417r-mediated induction of resistance. Furthermore, R fluorescens WCS417r-mediated systemic resistance did not coincide with the accumulation of PR mRNAs before challenge inoculation. These results indicate that R fluorescens WCS417r induces a pathway different from the one that controls classic systemic acquired resistance and that this pathway leads to a form of systemic resistance independent of SA accumulation and PR gene expression.

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