We utilized two approaches to probe the nature of gene complements within these two clades of oomycetes. First, selleck using the generalized approach of examining PANTHER protein families, we identi fied major lineage specific expansions of gene families. Second, through targeted analysis of subsets of the P. ultimum proteome, including the secretome, effectors, proteins involved in carbohydrate metabolism, and pathogen microbial associated molecular patterns. we revealed commonalities, as well as significant distinct features, of P. ultimum in comparison to Phytophthora spp. Over represented gene families Several families involved in proteolysis were over repre sented in P. ultimum compared to Phytophthora spp. This is primarily due to a massive expansion of subtilisin related proteases in P.
ulti mum following the divergence from ancestors of Phy tophthora. With regard to the total complement of serine proteases, the subtilisin family expansion in P. ultimum is somewhat counterbalanced Inhibitors,Modulators,Libraries by the tryp sin related serine protease family, which has undergone more gene duplication events in the Phytophthora line age than the Pythium lineage. The metalloprotease M12 family has also undergone multiple expansions, from one copy in the stramenopile most recent common ancestor, to three in the oomycete most recent common ancestor, then up to 12 in P. ultimum. E3 ligases are responsible for substrate specificity of ubiquitination and subsequent proteolysis, and secreted E3 ligases have been shown to act as effectors for patho gens by targeting host response proteins for degradation.
The HECT Inhibitors,Modulators,Libraries E3 family of ubiquitin protein ligases apparently underwent at least two major expansions, one in the oomycete lineage after the diver gence from diatoms and another in the P. ultimum line age. Most of the expansion in the P. ultimum lineage appears to be derived from repeated duplication of only two genes that were present in the Pythium Phytophthora common ancestor. This expanded subfamily is apparently ortholo gous to the UPL1 and UPL2 genes from A. thaliana. Inhibitors,Modulators,Libraries Of the 56 predicted HECT E3 ligases in the P. ultimum genome, 16 are predicted by SignalP to have bona fide signal peptides, and another 10 have Inhibitors,Modulators,Libraries predicted signal Inhibitors,Modulators,Libraries anchors, a substantially larger number than reported for other oomycete genomes. Under represented gene families Several gene families are significantly under represented in the P.
ultimum genome compared to Phytophthora and it appears that these are mostly due to expansions in the Phytophthora lineage rather than losses in the Pythium lineage, though the relatively long selleck products distance to the diatom outgroup makes this somewhat uncertain. These include the aquaporin family, the phospholipase D family, four families subfamilies of intracellu lar serine threonine protein kinases, and three families involved in sulfur metabolism, cysteine desulfurylases and sulfate trans porters. The P.