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The active enzyme in the oxidized state works as a homodimer linked by two intermolecular disulphide bonds between specific cysteines (Hothorn et al., 2006)

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In addition, some sHSPs, such as HSPB1/HSP27 and HSPB5, can be phosphorylated through kinase pathways, including the p38/MK2 pathway, which may regulate their chaperone activity, cytoskeletal interactions, and involvement in autophagy-related clearance of damaged cellular components (Ungelenk et al., 2016

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As ice crystals form, they can exert immense physical pressure on the peptide molecules, shearing and denaturing them