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To simplify, I will try to explain it as clearly as possible (refer to the illustration below for further clarification): During fermentation, protease enzymes in the dough break down gluten bonds, specifically breaking peptide bonds between proteins into shorter segments of amino acids

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The structural data obtained also provided a rationale for previously investigated mutations: Asp336, located at the C-terminus of the linker segment connecting GT1 and GT2 domains, whose hydrogen bond with GT2 Lys445 does not seem to be dramatically impacted when being mutated into a serine residue