Covalent intramolecular and intermolecular cross-links are formed in several steps. First, certain lysines and hydroxylysines are deaminated by the extracellular enzyme lysyl oxidase to yield highly reactive aldehyde groups. The aldehydes then react spontaneously to form covalent bonds with each other or with other lysines or hydroxylysines. Most of the cross-links form between the short nonhelical segments at each end of the collagen molecules.
From: The Extracellular Matrix of Animals
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