”’Heparan sulfate”’ (”’HS”’) is a linear [[polysaccharide]] found in all animal tissues.<ref>{{cite journal | vauthors = Medeiros GF, Mendes A, Castro RA, Baú EC, Nader HB, Dietrich CP | title = Distribution of sulfated glycosaminoglycans in the animal kingdom: widespread occurrence of heparin-like compounds in invertebrates | journal = Biochimica et Biophysica Acta (BBA) – General Subjects | volume = 1475 | issue = 3 | pages = 287–94 | date = July 2000 | pmid = 10913828 | doi = 10.1016/S0304-4165(00)00079-9 }}</ref> It occurs in a [[proteoglycan]] (HSPG, i.e. Heparan Sulfate ProteoGlycan) in which two or three HS chains are attached in close proximity to cell surface or [[extracellular matrix]] 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 MV | year=2000 | publisher=Marcel Dekker Inc. | location = New York, New York | pages=[https://archive.org/details/proteoglycansstr00iozz/page/n41 27]–59}}</ref><ref>{{cite journal | vauthors = Iozzo RV | title = Matrix proteoglycans: from molecular design to cellular function | journal = Annual Review of Biochemistry | volume = 67 | pages = 609–52 | year = 1998 | pmid = 9759499 | doi = 10.1146/annurev.biochem.67.1.609 | s2cid = 14638091 | doi-access = free }}</ref> In this form, HS binds to a variety of protein [[ligand]]s, including [[Wnt signaling pathway|Wnt]],<ref name=”Gao 576–587″>{{cite journal | vauthors = Gao W, Kim H, Feng M, Phung Y, Xavier CP, Rubin JS, Ho M | title = Inactivation of Wnt signaling by a human antibody that recognizes the heparan sulfate chains of glypican-3 for liver cancer therapy | journal = Hepatology | volume = 60 | issue = 2 | pages = 576–87 | date = August 2014 | pmid = 24492943 | pmc = 4083010 | doi = 10.1002/hep.26996 }}</ref><ref name=”Gao 26245″ /> and regulates a wide range of biological activities, including developmental processes, [[angiogenesis]], [[blood coagulation]], abolishing detachment activity by GrB ([[Granzyme B]]),<ref>{{cite journal | vauthors = Buzza MS, Zamurs L, Sun J, Bird CH, Smith AI, Trapani JA, Froelich CJ, Nice EC, Bird PI | display-authors = 6 | title = Extracellular matrix remodeling by human granzyme B via cleavage of vitronectin, fibronectin, and laminin | journal = The Journal of Biological Chemistry | volume = 280 | issue = 25 | pages = 23549–58 | date = June 2005 | pmid = 15843372 | doi = 10.1074/jbc.M412001200 | doi-access = free }}</ref> and tumour [[metastasis]]. HS has also been shown to serve as cellular receptor for a number of viruses, including the [[respiratory syncytial virus]].<ref>{{cite journal | vauthors = Hallak LK, Spillmann D, Collins PL, Peeples ME | title = Glycosaminoglycan sulfation requirements for respiratory syncytial virus infection | journal = Journal of Virology | volume = 74 | issue = 22 | pages = 10508–13 | date = November 2000 | pmid = 11044095 | pmc = 110925 | doi = 10.1128/JVI.74.22.10508-10513.2000 }}</ref> One study suggests that cellular heparan sulfate has a role in SARS-CoV-2 infection, particularly when the virus attaches with ACE2.<ref>{{cite journal | vauthors = Clausen TM, Sandoval DR, Spliid CB, Pihl J, Perrett HR, Painter CD, Narayanan A, Majowicz SA, Kwong EM, McVicar RN, Thacker BE | display-authors = 6 | title = SARS-CoV-2 Infection Depends on Cellular Heparan Sulfate and ACE2 | journal = The Journal of Cell|date = 14 September 2020| volume = 183 | issue = 4 | pages = 1043–1057.e15 |pmid = 32970989 | doi = 10.1016/j.cell.2020.09.033 | pmc = 7489987 }}</ref>
”’Heparan sulfate”’ (”’HS”’) is a linear [[polysaccharide]] found in all animal tissues.<ref>{{cite journal | vauthors = Medeiros GF, Mendes A, Castro RA, Baú EC, Nader HB, Dietrich CP | title = Distribution of sulfated glycosaminoglycans in the animal kingdom: widespread occurrence of heparin-like compounds in invertebrates | journal = Biochimica et Biophysica Acta (BBA) – General Subjects | volume = 1475 | issue = 3 | pages = 287–94 | date = July 2000 | pmid = 10913828 | doi = 10.1016/S0304-4165(00)00079-9 }}</ref> It occurs in a [[proteoglycan]] (HSPG, i.e. Heparan Sulfate ProteoGlycan) in which two or three HS chains are attached in close proximity to cell surface or [[extracellular matrix]] 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 MV | year=2000 | publisher=Marcel Dekker Inc. | location = New York, New York | pages=[https://archive.org/details/proteoglycansstr00iozz/page/n41 27]–59}}</ref><ref>{{cite journal | vauthors = Iozzo RV | title = Matrix proteoglycans: from molecular design to cellular function | journal = Annual Review of Biochemistry | volume = 67 | pages = 609–52 | year = 1998 | pmid = 9759499 | doi = 10.1146/annurev.biochem.67.1.609 | s2cid = 14638091 | doi-access = free }}</ref> In this form, HS binds to a variety of protein [[ligand]]s, including [[Wnt signaling pathway|Wnt]],<ref name=”Gao 576–587″>{{cite journal | vauthors = Gao W, Kim H, Feng M, Phung Y, Xavier CP, Rubin JS, Ho M | title = Inactivation of Wnt signaling by a human antibody that recognizes the heparan sulfate chains of glypican-3 for liver cancer therapy | journal = Hepatology | volume = 60 | issue = 2 | pages = 576–87 | date = August 2014 | pmid = 24492943 | pmc = 4083010 | doi = 10.1002/hep.26996 }}</ref><ref name=”Gao 26245″ /> and regulates a wide range of biological activities, including developmental processes, [[angiogenesis]], [[blood coagulation]], abolishing detachment activity by GrB ([[Granzyme B]]),<ref>{{cite journal | vauthors = Buzza MS, Zamurs L, Sun J, Bird CH, Smith AI, Trapani JA, Froelich CJ, Nice EC, Bird PI | display-authors = 6 | title = Extracellular matrix remodeling by human granzyme B via cleavage of vitronectin, fibronectin, and laminin | journal = The Journal of Biological Chemistry | volume = 280 | issue = 25 | pages = 23549–58 | date = June 2005 | pmid = 15843372 | doi = 10.1074/jbc.M412001200 | doi-access = free }}</ref> and tumour [[metastasis]]. HS has also been shown to serve as cellular receptor for a number of viruses, including the [[respiratory syncytial virus]].<ref>{{cite journal | vauthors = Hallak LK, Spillmann D, Collins PL, Peeples ME | title = Glycosaminoglycan sulfation requirements for respiratory syncytial virus infection | journal = Journal of Virology | volume = 74 | issue = 22 | pages = 10508–13 | date = November 2000 | pmid = 11044095 | pmc = 110925 | doi = 10.1128/JVI.74.22.10508-10513.2000 }}</ref> One study suggests that cellular heparan sulfate has a role in SARS-CoV-2 infection, particularly when the virus attaches with ACE2.<ref>{{cite journal | vauthors = Clausen TM, Sandoval DR, Spliid CB, Pihl J, Perrett HR, Painter CD, Narayanan A, Majowicz SA, Kwong EM, McVicar RN, Thacker BE | display-authors = 6 | title = SARS-CoV-2 Infection Depends on Cellular Heparan Sulfate and ACE2 | journal = The Journal of Cell|date = 14 September 2020| volume = 183 | issue = 4 | pages = 1043–1057.e15 |pmid = 32970989 | doi = 10.1016/j.cell.2020.09.033 | pmc = 7489987 }}</ref>
