The Hrp pathogenicity island of Erwinia amylovora and identification of three novel genes required for systemic infection

Sequence analysis of the region bordering the hrp/dsp gene cluster of Erwinia amylovora strain Ea321, which causes fire blight, revealed characteristics of pathogenicity islands (PAIs). Included are genes for a phage integrase, a tRNA^Phe, several orthologues of genes of YAPI, a PAI of Yersinia pseudotuberculosis, and several putative virulence genes with HrpL-dependent promoter motifs. The island is designated the Hrp PAI of E. amylovora. It is comprised of a chromosomal region of c. 62 kb with 60 open reading frames (ORFs). Comparison of the Hrp PAI of E. amylovora with those of four closely related bacteria showed that orfB, a homologue of avrBsT of Xanthomonas campestris pv. vesicatoria, and orfA, its putative chaperone gene, are present only in the Hrp PAI of E. amylovora. As regions flanking the hrp/dsp gene cluster are quite diverse, addition and deletion may have occurred during divergent evolution of the five bacteria. Among ORFs of the PAI of Ea321, three new HrpL-dependent genes were identified. Because they are required for full virulence in apple, they were designated hsvC, hsvB and hsvA (hrp-associated systemic virulence). They encode a homologue of an amidinotransferase for phaseolotoxin biosynthesis and homologues of a nikkomycin-biosynthetic protein of Pseudomonas syringae.