Mouse protein 25 (Mo25) stimulates Fray activity in vitro, and it is required for tubule transepithelial transport in stimulated (hypotonic) conditions. (A) Mo25 stimulates autophosphorylation of Fray (Drosophila Ste20-related proline/alanine-rich kinase [SPAK]/oxidative stress response 1 [OSR1] homolog; upper two bands) as well as sodium-potassium 2 chloride cotransporter (NKCC) phosphorylation (lower band) in vitro. The effect is more pronounced with FrayT206E, in which the T-loop Thr that is the target of With No Lysine kinase (WNK) phosphorylation is mutated to a phosphomimicking Glu. In vitro kinase assays were performed with purified Drosophila Fray, D. melanogaster mouse protein 25 (DmMo25), and the N terminus of Drosophila NKCC, Ncc69 (amino acids 1–204). A representative gel is shown of three independent experiments (Supplemental Figure 5). WT, wild type. (B) DmMo25 is required in the tubule principal cell for hypotonicity-stimulated transepithelial potassium flux. Transepithelial potassium flux was measured in control tubules (w; c42-GAL4/+) and DmMo25 knockdown tubules (w; UAS-DmMo25RNA interference (RNAi)/+; c42-GAL4/+) bathed in isotonic standard bathing medium or hypotonic medium (standard bathing medium diluted with water). Two-way ANOVA revealed significant effects of genotype (P<0.001), condition (P<0.001), and interaction (P<0.01). In a Sidak multiple comparisons test, hypotonic conditions resulted in a significant increase in potassium flux in control tubules but no change in DmMo25 knockdown tubules (P=0.82). There was no significant difference between control and DmMo25 knockdown tubules in isotonic conditions (P=0.67), whereas a significant difference was observed in hypotonic conditions (P<0.001). ****Adjusted P value <0.001. (C) Expression of mammalian Mo25 rescues the decreased potassium flux phenotype of DmMo25 knockdown tubules. Transepithelial potassium flux was measured in hypotonic conditions in control tubules (w; c42-GAL4/+), DmMo25 principal cell knockdown tubules (w; UAS-DmMo25RNAi/+; c42-GAL4/+), and DmMo25 knockdown tubules expressing mouse Mo25 (w; UAS-DmMo25RNAi/+; UAS-MmMo25/c42-GAL4). One-way ANOVA (P<0.001). In a Tukey multiple comparison test comparing all genotypes with one another, adjusted P values were <0.001 for the difference between w; c42-GAL4/+ and w; UAS-DmMo25RNAi/+; c42-GAL4/+ and also for the difference between w; UAS-DmMo25RNAi/+; c42-GAL4/+ and w; UAS-DmMo25RNAi/+; UAS-MmMo25/c42. Adjusted P value was 0.04 for the difference between w; c42-GAL4/+ and w; UAS-DmMo25RNAi/+; UAS-MmMo25/c42-GAL4. ****Adjusted P value <0.001.