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Table 3 Amino acids in collagen with their corresponding post-translation modification, added group and enzyme involved. The outcome of overexpression of each modification is described in the last column

From: Understanding the matrix: collagen modifications in tumors and their implications for immunotherapy

Amino acids

PTM

Group

Enzymes

Outcome if increased

Proline

Hydroxylation (ER)

–OH

P4Hs, P3Hs

Hypoxia

Lysine

Hydroxylation (ER)

–OH

LH1, LH2, LH3

Increased glycosylation, crosslinking and fibrosis

Hydroxylysine

N-Linked (ER)

O-linked (GA) Glycosylation

Glucose or galactose

COLGALT1, COLGALT2,

GT, GGT

Increased crosslinking, matrix stiffness

Lysine/Hydroxylysine

Oxidation (ES)

–O

LOX, LOXL1, LOXL2, LOXL3, LOXL4

Fibrosis, matrix stiffness

Arginine

Citrullination (ES)

Citrulline

H2O to NH4 + 

PAD4

Role in protein folding, apoptosis, TGF-β

Pathway, receptor binding