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GSH biosynthesis occurs exclusively in the cytosol via a two-step ATP-dependent reaction catalysed sequentially by two enzymes: gamma-glutamylcysteine synthetase (GCL also referred to as gamma-glutamylcysteine ligase, comprising the catalytic GCLC subunit and regulatory GCLM subunit), which catalyses the rate-limiting first step of joining glutamate and cysteine