The hrp gene cluster from Xanthomonas campestris pv. vesicatoria determines functions necessary not only for pathogenicity on the host plants pepper and tomato but also for the elicitation of the hypersensitive reaction on resistant host and nonhost plants. Transcriptional orientation and expression of the hrp loci were determined with hrp::Tn3-gus fusions. In addition, expression of the hip loci was studied by RNA hybridization experiments. Expression of the hrp genes was not detectable after growth of the bacteria in complex medium or in minimal medium. However, high levels of induction of hrp gene expression were measured during growth of the bacteria in the plant. To search for a plant molecule responsible for this induction, we examined a variety of materials of plant origin for their ability to induce hrp gene expression. Filtrates from plant suspension cultures induced hrp genes to levels comparable to those induced in the plant. The inducing molecule(s) was found to be heat stable and hydrophilic and to have a molecular mass of less than 1,000 daltons.The molecular mechanisms involved in plant-bacterium interactions during pathogenesis are complex and far from being understood. In the last few years, a number of bacterial genes that determine the outcome of the interaction between the bacterium and the plant have been identified and isolated. Most notable are two classes of genes required for basic compatibility: disease-specific (dsp) genes, which are associated with disease development in host plants but not with the induction of a hypersensitive response (HR) in nonhost plants (7, 27); and hrp genes, which are required for both the pathogenic interaction with host plants and the induction of the HR in resistant host and nonhost plants. hrp genes have been cloned from a number of different species of gram-negative phytopathogenic bacteria, e.g., Erwinia amylovora, Pseudomonas solanacearum, and pathovars of Pseudomonas syringae and ofXanthomonas campestris (for a review, see reference 35). Genetic analysis of hrp genes from these different organisms indicates that they determine basic pathogenicity functions necessary for any interaction with the plant. The elucidation of their biochemical function and their role in the plant-bacterium interaction is expected to lead to a molecular understanding of the mechanisms underlying bacterial plant diseases.We have chosen the interaction between X. campestris pv. vesicatoria, the causal agent of bacterial spot disease, and its host plants, pepper (Capsicum annuum L.) and tomato (Lycopersicum esculentum Mill.), as a system for the analysis of hrp genes. After invasion of the plant via stomata or wounds, X. campestris pv. vesicatoria multiplies in the intercellular spaces of the leaf tissue, giving rise to disease symptoms (29). Depending on the susceptibility of the particular plant cultivar, two different types of reactions can be observed. In the susceptible plant, water-soaked lesions occur (compatible interaction). In the resistant plant, avirulent strains in...