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MBE Advance Access published online on October 19, 2005
Molecular Biology and Evolution, doi:10.1093/molbev/msj039
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© The Author 2005. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oxfordjournals.org
Accepted October 10, 2005

Research Article

Ancient Origin of Elicitin Gene Clusters in Phytophthora Genomes

Rays H.Y. Jiang 1, Brett M. Tyler 2, Stephen C. Whisson 3, Adrienne R. Hardham 4, and Francine Govers 1*

1 Laboratory of Phytopathology, Wageningen University, Binnenhaven 5, NL-6709 PD, Wageningen and Graduate School Experimental Plant Sciences, The Netherlands
2 Virginia Bioinformatics Institute, Virginia Polytechnic and State University, Blacksburg VA, USA
3 Plant Pathogen Interactions Programme, Scottish Crop Research Institute, Invergowrie, Dundee DD2 5DA, Scotland, UK
4 Plant Cell Biology Group, Research School of Biological Sciences, Australian National University, Canberra, ACT 2601, Australia

* To whom correspondence should be addressed.
Francine Govers, E-mail: Francine.Govers{at}wur.nl


  Abstract

The genus Phytophthora belongs to the oomycetes in the eukaryotic stramenopile lineage and is comprised of over 65 species that are all destructive plant pathogens on a wide range of dicotylodons. Phytophthora produces elicitins (ELIs), a group of extracellular elicitor proteins that cause a hypersensitive response in tobacco. Database mining revealed several new classes of elicitin-like (ELL) sequences with diverse elicitin domains in Phytophthora infestans, Phytophthora sojae, Phytophthora brassicae and Phytophthora ramorum. ELIs and ELLs were shown to be unique to Phytophthora and Pythium species. They are ubiquitous among Phytophthora species and belong to one of the most highly conserved and complex protein families in the Phytophthora genus. Phylogeny construction with elicitin domains derived from 156 ELIs and ELLs showed that most of the diversified family members existed prior to divergence of Phytophthora species from a common ancestor. Analysis to discriminate diversifying and purifying selection showed that all 17 ELI and ELL clades are under purifying selection. Within highly similar ELI groups there was no evidence for positively selected amino acids suggesting that purifying selection contributes to the continued existence of this diverse protein family. Characteristic cysteine spacing patterns were found for each phylogenetic clade. Except for the canonical clade ELI-1, ELIs and ELLs possess C-terminal domains of variable length, many of which have a high threonine, serine or proline content suggesting an association with the cell wall. In addition, some ELIs and ELLs have a predicted GPI (glycosylphosphatidylinositol) site suggesting anchoring of the C-terminal domain to the cell membrane. The eli and ell genes belonging to different clades are clustered in the genomes. Overall, eli and ell genes are expressed at different levels and in different life cycle stages but those sharing the same phylogenetic clade appear to have similar expression patterns.

Keywords: elicitin; Phytophthora; molecular phylogeny.
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