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The second step in GSH synthesis is catalyzed by glutathione synthetase (encoded by GSHS ) and involves the addition of glycine to -Glu-Cys ( hGSHS gene ( cis -regulatory elements are present in the promoter regions of both GSHS and hGSHS , indicating that, in legumes, they may either perform different functions or perform similar functions that are separated temporally or spatially ( Medicago sativa , GSH is associated with meristematic cells, cell-cycle activation, and the induction of somatic embryogenesis, while hGSH is associated with differentiated cells and embryo proliferation ( Lotus japonicus , meanwhile, GSHS is expressed only in nodules, whereas hGSHS is also expressed in leaves and roots ( M