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Extracellular matrix
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====Heparan sulfate==== [[Heparan sulfate]] (HS) is a linear [[polysaccharide]] found in all animal tissues. It occurs as a [[proteoglycan]] (PG) in which two or three HS chains are attached in close proximity to cell surface or ECM proteins.<ref>{{cite book | title=Proteoglycans: structure, biology and molecular interactions | url=https://archive.org/details/proteoglycansstr00iozz | url-access=limited | vauthors = Gallagher JT, Lyon M | chapter=Molecular structure of Heparan Sulfate and interactions with growth factors and morphogens | veditors = Iozzo RV | year=2000 | publisher=Marcel Dekker Inc. New York, New York | pages=[https://archive.org/details/proteoglycansstr00iozz/page/n41 27]β59 |isbn=9780824703349 }}</ref><ref>{{cite journal | vauthors = Iozzo RV | s2cid = 14638091 | title = Matrix proteoglycans: from molecular design to cellular function | journal = Annual Review of Biochemistry | volume = 67 | issue = 1 | pages = 609β52 | year = 1998 | pmid = 9759499 | doi = 10.1146/annurev.biochem.67.1.609 | doi-access = free }}{{closed access}}</ref> It is in this form that HS binds to a variety of protein [[ligand]]s and regulates a wide variety of biological activities, including [[developmental processes]], [[angiogenesis]], [[blood coagulation]], and tumour [[metastasis]].{{cn|date=April 2025}} In the extracellular matrix, especially [[basement membrane]]s, the [[protein domain|multi-domain]] proteins [[perlecan]], [[agrin]], and [[type XVIII collagen|collagen XVIII]] are the main proteins to which heparan sulfate is attached.{{cn|date=April 2025}}
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