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Following isolation from G. rostochiensis and H. glycines [ 5 ] , cellulases have been identified in M. incognita [ 6 ] , G. tabacum [ 68 ] , H. schachtii [ 69 ] , and P. penetrans [ 70 ] . Additional prokaryotic-like sequences identified in plant parasitic nematodes include other cell-wall-degrading enzymes such as xylanase [ 7 ] , pectate lyase [ 8 71 ] and polygalacturonase [ 72 ] , and evidence is accumulating that these sequences have been acquired by horizontal gene transfer [ 11 ] . The known Meloidogyne cellulase (MI00483.cl), potentially novel cellulases (MI00537.cl, MI01196.cl, MI01381.cl, MI01842.cl), and pectate lyase (MI00592.cl, MI00520.cl) were represented in the M. incognita EST clusters.
On the basis of biochemical and immunological criteria, genes have been identified in Globodera rostochiensis and Heterodera glycines that allow these nematodes to endogenously produce enzymes that can degrade cellulose and pectin, the two major components of plant cell walls.
The pectate lyase gene (MI00592) was identified in G. rostochiensis and H. glycines [ 17 ] and the exo-polygalacturonase (MI00252) was identified in our M. incognita data [ 21 22 ] . Because of the obvious role of nematode genes that allow endogenous production of cell-wall degrading enzymes in attacking a plant host, it has been hypothesized that their acquisition by HGT may have been key steps in the evolution of plant-parasitic nematodes from ancestral free-living forms [ 3 ] . In that model, an intermediate, symbiotic association of a soil-dwelling (but free-living) nematode with a soil bacterium possessing these enzymes is postulated before the HGT event.
Use of SL1 by transcripts is estimated at 70% in C. elegans [ 60 ] , more than 80% in Ascaris lumbricoides [ 61 ] , and approximately 60% in G. rostochiensis (Ling Qin, personal communication).
Another RKN gene also postulated to have been acquired by HGT, and which encodes polyglutamate synthetase, was previously identified in M. artiellia [ 23 ] . Significantly, hybridization data showed that this particular gene is absent from both the M. javanica and G. rostochiensis genomes [ 23 ] . We speculate that acquisition of this gene by M. artiellia is a recent HGT event (event 'c', Figure 1), and thus it is truly absent from the Meloidogyne genomes from which our datasets were derived.
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