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According to Korkushko et al

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The constructed pURA3-GSH1 strain was heterozygous at the GSH1 locus, with one allele expressing the URA3 ORF under pGSH1 control and the other expressing GSH1 under pTEF1 control (FigureS1 A )

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The release of vasoactive intestinal peptide (VIP) and acetylcholine (Ach) by submucosal neurons stimulates the enterocyte and Cl enters by co-transport with sodium and potassium through the basolateral Na + -K + -2Cl channel