SUMMARY The genus Botrytis contains necrotrophic plant pathogens that have a wide host range (B. cinerea) or are specialized on a single host species, e.g. B. elliptica on lily. In this study, it was found that B. elliptica-induced cell death of lily displays hallmark features of animal programmed cell death or apoptosis including cytoplasmic shrinkage, nuclear DNA fragmentation and the accumulation of NO as well as H(2)O(2). A pharmacological approach showed that B. elliptica-induced cell death could be modulated by serine and cysteine protease inhibitors including one caspase inhibitor. Blocking phosphatase activity stimulated cell death and concomitant lesion formation, suggesting that B. elliptica-induced cell death is mediated by kinase/phosphatase pathways. Blocking Ca(2+) influx restricted cell death. Blocking steps of sphingolipid biosynthesis delayed lily cell death for several days. B. elliptica culture filtrate (CF) was able to induce lily cell death by means of secreted proteins. Induction of cell death is necessary and sufficient for pathogenicity and host specialization because prior infiltration of B. elliptica CF enabled subsequent infection of lily by the otherwise incompatible pathogens B. cinerea and B. tulipae. The secreted B. elliptica proteins also induced cell death in some but not all Arabidopsis accessions and mutants. Arabidopsis accessions that respond to infiltration of B. elliptica CF also display cell death symptoms upon inoculation with B. elliptica conidia.