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Population Doubling Level Calculation To calculate the PDL, a formula was used as follows: n=3.32 (log UCY log I)+X, where nis the nal PDL at the end of a given subculture, UCY is the cell yield at that point, Iis the cell number used as inoculum to begin that subculture, and Xis the PDL of the inoculum used to initiate the subculture being quantitated (Pham et al., 2020)