, 2011, Su et al., 2012, DeLay et al., 2012 and Ji et al., 2013) (Table S1). In the GRH salivary transcriptome, three unique contigs of endo-beta glucanase (EGase) (comp12770, comp10542, and comp96112)

(EC 3.2.1.4), and one contig of alpha amylase (comp218776) (EC 3.2.1.1) were predicted, although their expression levels were not high (Table S1). EGase cleaves amorphous sites of cellulose chains (Watanabe and Tokuda, 2010), and xyloglucan. Alpha amylase hydrolyzes the bonds of oligosaccharides and polysaccharides. Cellulose and polysaccharides are major components of plant cell walls, and xyloglucan is abundant in phloem cell walls MK-1775 manufacturer of commelinid monocotyledons, including rice plants (Brennan and Harris, 2011 and Yokoyama Fulvestrant mouse and Nishitani, 2014). In the E. fabae sialotranscriptome, 58 EGases and 36 alpha amylases ( DeLay et al., 2012), and in BPH, one EGase,

two beta-1,3-glucanases, and one alpha amylase were found ( Ji et al., 2013). In B. tabaci, two alpha amylases were predicted in the transcriptome ( Su et al., 2012). They are putative degrading enzymes for plant cell walls, facilitating sap-sucking. Among detoxification-associated proteins, P450 and GST are expected to be important for resistance to xenobiotics including plant allelochemicals and insecticides (Feyereisen, 2005, Després et al., 2007 and Li et al., 2007). GRH contained 59 putative P450s and 20 glutathione-S-transferases (GSTs) as unique contigs,

with various expression levels (Table S1). In BPH, 63 ROS1 P450s and one GST were found (Ji et al., 2013). In E. fabae, 41 P450s and four GSTs were predicted ( DeLay et al., 2012), in contrast to eight and five, respectively, in B. tabaci, ( Su et al., 2012) and only one and three in A. pisum ( Carolan et al., 2011). Aphids and whiteflies including A. pisum and B. tabaci penetrate the epidermis with their stylets and intercellularly probe parenchyma cells before reaching the phloem ( Nault and Gyrisco, 1966, Walker and Perring, 1994 and Jiang et al., 1999), although brief cell punctures are performed during intercellular penetration ( Tjallingii, 1985 and Janssen et al., 1989). In contrast, GRH and BPH intracellularly penetrate mesophyll or parenchyma cells until the vascular bundles are encountered ( Naito and Masaki, 1967 and Sogawa, 1982). E. fabae is both a cell-rupturing and sheath feeder and mechanically injures parenchyma cells and phloem ( Backus et al., 2005). Thus, these Auchenorrhyncha species puncture the parenchyma or mesophyll cells, thereby come into contact with defensive chemicals that are stored within vacuoles and apoplasts. This behavior may explain why GRH and other hoppers possess multiple P450 (isoforms), which catalyze the oxidation of diverse secondary substances.