, 2005). Both genomes also encode proteins (GI:289669426 and GI:289663837) sharing Roxadustat order 30% amino acid sequence identity with the putative T3SS effector RipT (RSc3212), a YopT-like cysteine protease from the betaproteobacterium Ralstonia solanacearum GMI1000 (Poueymiro & Genin, 2009). Close homologues are not found in any other Xanthomonas genomes, but a protein (GI:270492983) from another plant-pathogenic betaproteobacterium, Acidovorax avenae ssp. avenae ATCC 19860, shares 48% sequence identity with the Xvv and Xcm RipT-like proteins. There are some differences between Xcm 4381 and Xvv 702 with respect to their complements of effectors that might contribute to their different host ranges. Xcm 4381 encodes two predicted

YopJ-like C55 cysteine proteases (GI:289670655 and GI:289671144) that are absent from Xvv 702. On the other hand, Xvv 702 encodes a protein (GI:289661936) sharing 87% amino acid sequence identity with Xanthomonas euvesicatoria XopAF (also known as AvrXv3) (Astua-Monge et al., 2000). This gene is absent from Xcm 4381, but shares 35% identity (at the amino acid level) with the HopAF1-like genes found at the integron locus in both Xcm and Xvv. Such differences in effector repertoires

have previously been shown to be significant for host adaptation (Wei et al., 2007; Kvitko et al., 2009; SB203580 in vitro Lindeberg et al., 2009). For example, HopQ1-1 is present in P. syringae pathovar phaseolicola, where it suppresses immunity in beans, but is absent from P. syringae pathovar tabaci, and triggers defences in tobacco (Ferrante et al., 2009). It is possible Pregnenolone that the differences in effector repertoires of Xcm 4381 and Xvv 702 are significant

for the adaptation of Xcm 4381 to a new host (i.e. banana). It remains to be tested whether the two Xcm 4381 YopJ- and HopR-like proteins suppress defences and whether the Xvv 702 AvrXv3 confers avirulence in banana. The outer membranes of Gram-negative bacteria are covered with lipopolysaccharides (Lerouge & Vanderleyden, 2002). Among different strains of X. campestris pathovar campestris and X. oryzae pathovar oryzae, the lipopolysaccharide biosynthesis locus shows hypervariability arising from horizontal transfer (Patil & Sonti, 2004; Patil et al., 2007). The lipopolysaccharide locus in Xcm 4381 (GenBank: ACHT01000245.1) most closely matches that of Xanthomonas axonopodis pathovar citri 306 (93% nucleotide sequence identity). The lipopolysaccharide locus in Xvv 702 (GenBank: ACHS01000380.1) shows no significant sequence similarity to that of Xcm 4381. It does, however, share 86% nucleotide sequence identity with Xanthomonas albilineans strain GPE PC73 (Pieretti et al., 2009). This is incongruent with the close phylogenetic relationship between Xcm 4381 and Xvv 702 and indicates recent horizontal transfer in one or both strains from independent sources. Any significance of this variation between Xcm 4381 and Xvv 702 for virulence and host specificity remains unclear.