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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.
Sixty-four ESTs had significant homology to the putative fatty-acid-binding protein Sec-2, confirming the abundant expression of this gene reported in L2 cDNA libraries from M. incognita [ 29 ] and the cyst nematodes G. rostochiensis and G. pallida cDNA [ 30 ] . Sec-2 is secreted by plant-parasitic nematodes at relatively high levels [ 34 ] . Several abundantly expressed genes are also horizontal gene transfer candidates (see below).
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.
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.
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).
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