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The substitutions are at positions 2 (D-Ala for Ala — protects the N-terminus from dipeptidyl peptidase IV), position 8 (Gln for Asn — reduces deamidation), position 15 (Ala for Gly — reduces aspartic-acid attack at this site), and position 27 (Leu for Met — prevents methionine oxidation)