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found that lysosomal PEN2 is recruited to ATP6AP1 subunit upon binding to metformin to regulate v-ATPase, which provides the impetus for the translocation of AXIN and liver kinase B1 (LKB1) to the lysosomal surface to activate AMPK, enabling low concentrations of metformin to utilize the AMP-independent AMPK activation pathway (8)

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