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. 2021 Dec 22;23(1):91. doi: 10.3390/ijms23010091

Table 1.

The enzymes involved in the crucial stages of collagen formation.

Intracellular Stage of
Collagen Formation
Enzyme Gene miRNA References
1. COL1A1 gene expression COL1A1 miR-625-5p,
miR-92a-1-5p,
miR-129-5p,
miR-133b,
miR-133a-3p,
miR-196b-5p,
miR-196a,
miR-98,
miR-29a,
miR-513b-5p
[12,13,15,16,17,18,23,24,25,26,27,28,29]
2. Hydroxylation of proline residues Prolyl,4- hydroxylase P4HA1,

P4HA2, P4HA3,
P4HB
miR-124-3p,
miR-122,
miR-30e,

miR-210
[30,31,32]
[33,34]
3. Hydroxylation of lysine residues Lysyl hydroxylase PLOD1,
PLOD2,

PLOD3
miR-34c,
miR-124,
miR-26,
miR-663a
[35,36,37,38]
4. Glycosylation of some hydroxylysine residues Glycosyltransferase GLT25D1
GLT25D2
[39,40,41]
5. Formation of intra-chain and inter-chain disulphide bonds in terminal peptides Protein disulphide isomerase P4HB miR-210 [33,34]
6. Procollagen triple helix formation Hsp-47 protein SERPINH1 miR-29b,
miR-29a
[42,43,44]
Extracellular stage of collagen formation Enzyme Gene miRNA Sources
7. Cleavage of amino propeptides Specific N-propeptidases ADAM2,3 and 14 miR-29b [45]
8. Cleavage of carboxy propeptides Specific C- propeptidase BMP1 miR-194,
miR-29c,
miR-29b
[45,46,47]
9. Aggregation of collagen fibres Self-assembly [48]
10. Crosslinking
Oxidative deamination of epsilon-amino groups of lysyl and hydroxylysine residues located in telopeptide domains to aldehydes
Lysyl oxidase LOX miR-27,
miR29b,
miR29a,
miR-30a,
miR-30b,
miR-30b-2-3p,
miR-142-3p
[43,49,50,51,52,53,54,55]