Several encystation-specific genes have been identified and characterized

during the last decade, and have shown to be up-regulated with similar kinetics during encystation, suggesting that their regulation is at the transcriptional level [70]. Several reports also described putative transcription factors that regulate the expression of encystation-specific genes [71–74]. It was assumed that the encystation process is controlled at multiple levels (basic transcription, enhancement or de-repression) [62]. Moreover, it was hypothesized that epigenetic chromatin modifications via histone acetylation/deacetylation may participate in modulation of stage differentiation in this parasite [75]. In higher organisms, different RNA helicases have been described to interact with histone deacetylases (HDACs), such PF-562271 manufacturer as the known transcriptional regulator DP103 (Ddx20, Gemin3), which was found to immunoprecipitate with histone deacetylases HDAC2 and HDAC5, suggesting a role in transcription repression through HDACs recruitment [76]. In addition, the role of the RNA helicases p68 (Ddx5) and p72 (Ddx17) as transcription repressors when interacting with HDAC1 [77], HDAC2 and HDAC3 has been reported [78]. Our findings regarding the levels

of induction of the RNA helicase genes by qPCR were diverse, LB-100 purchase ranging from a smooth 2-4-fold induction in some DEAD-box genes to a high (20-31 times) relative expression in other genes.

Two genes, DEAD-box GL50803_13791 and DEAH-box GL50803_13200, presented a marked induction of 554 and 228 times, respectively, under the encystation conditions. Notably, the up-regulation of the encystation-specific gene coding for CWP2 increased up to 2,187 times compared to its expression in trophozoites. In Giardia, the RNAi machinery controlling antigenic variation has been found to involve a Dicer Galeterone enzyme with unique characteristics when compared to Dicer enzymes from higher eukaryotes. Giardia Dicer lacks the DExD/H helicase domain as well as double-stranded RNA binding motifs present in other Dicer homologs. Because we are only starting to understand the different roles of RNA helicases in RNAi, there are still many unresolved questions. Since different RNA helicases might operate at different steps in the RNAi pathway or might play different roles, the presence of thirty two putative DExD/H-box helicases in the Giardia genome and their differential patterns of expression during antigenic variation PF-4708671 order support their importance for RNAi. It would be relevant to determine the role of particular Giardia RNA helicases for different subsets of miRNA or siRNAs.