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This is best seen in examining the expression Parvulin of the Ribi genes (Figure?2B). Here removal of all PP-IPs, by deletion of Kcs1 and Vip1, causes the largest defect in gene repression. However, deletion of Arg82, which blocks production of 1-PP-IP5, 5-PP-IP4 and 5-PP-IP5 via pyrophosphorylation of IP5 and IP6 (but allows some production of PP-IP3 and PP-IP4 by Kcs1 dependent pyrophosphorylation of IP3 (Seeds et?al., 2005) also has a dramatic defect in gene repression. Finally, deletion of Ipk1 also causes a defect in Ribi gene repression, but less than that found in kcs1��vip1�� or arg82�� cells, as expected since ipk1�� cells can still synthesize 5-PP-IP5. Overall, therefore, the defect in Ribi gene repression in stress is proportional to the number of PP-IPs lost in a strain: kcs1��vip1��?BEZ235 cell line VE 821 By contrast, examining the influence of the PP-IP pathway on the induction of stress genes reveals two behaviors. Some genes show the trend described above for PP-IP dependent gene repression, where expression is proportional to the number of PP-IPs removed ( Figure?S5C). However, the majority of genes only have a significant expression defect when both Kcs1 and Vip1 are deleted ( Figure?S5D). Therefore, PP-IP dependent gene activation is generally less sensitive to a decrease in PP-IP level than PP-IP dependent gene repression. This is entirely consistent with our model where PP-IPs activate Rpd3L, which then acts as both an activator and repressor ( Figure?4E), since gene repression and gene activation by Rpd3L are likely to occur via different mechanisms and thus have different dependencies on PP-IP concentration. As discussed in the main text of the paper Watson et?al. found that HDAC3 is activated by IP4. IP4 acts by promoting the interaction between HDAC3 and the Corepressor SMRT. Although SMRT has a homolog in yeast Snt1, this protein does not play a role in the ESR as a snt1�� cells have no change in gene expression in YEPD or KCl stress conditions ( Table S1). Over the last 5 years, the PP-IPs have emerged as important signaling molecules in eukaryotic cells.