Gene Symbol: COL4A1
Description: collagen type IV alpha 1 chain
Alias: BSVD, RATOR, collagen alpha-1(IV) chain, COL4A1 NC1 domain, arresten, collagen IV, alpha-1 polypeptide, collagen of basement membrane, alpha-1 chain
Species: human
Products:     COL4A1

Top Publications

  1. Gould D, Phalan F, Breedveld G, van Mil S, Smith R, Schimenti J, et al. Mutations in Col4a1 cause perinatal cerebral hemorrhage and porencephaly. Science. 2005;308:1167-71 pubmed
    ..We show that vascular defects are caused by a semidominant mutation in the procollagen type IV alpha 1 gene (Col4a1) in mice, which inhibits the secretion of mutant and normal type IV collagen...
  2. Plaisier E, Chen Z, Gekeler F, Benhassine S, Dahan K, Marro B, et al. Novel COL4A1 mutations associated with HANAC syndrome: a role for the triple helical CB3[IV] domain. Am J Med Genet A. 2010;152A:2550-5 pubmed publisher
    The COL4A1 gene encodes the ?1-chain of type IV collagen, which is ubiquitously expressed in basement membranes...
  3. Plaisier E, Gribouval O, Alamowitch S, Mougenot B, Prost C, Verpont M, et al. COL4A1 mutations and hereditary angiopathy, nephropathy, aneurysms, and muscle cramps. N Engl J Med. 2007;357:2687-95 pubmed
    ..Linkage studies involving microsatellite markers flanking the COL4A1-COL4A2 locus were performed, followed by sequence analysis of COL4A1 complementary DNA extracted from skin-..
  4. Livingston J, Doherty D, Orcesi S, Tonduti D, Piechiecchio A, La Piana R, et al. COL4A1 mutations associated with a characteristic pattern of intracranial calcification. Neuropediatrics. 2011;42:227-33 pubmed publisher of ICC, we identified 5 patients with characteristic radiological features, in whom a mutation in the COL4A1 gene was found.All patients had CT and MR imaging...
  5. Rødahl E, Knappskog P, Majewski J, Johansson S, Telstad W, Krakenes J, et al. Variants of anterior segment dysgenesis and cerebral involvement in a large family with a novel COL4A1 mutation. Am J Ophthalmol. 2013;155:946-53 pubmed publisher
    ..revealed that they shared a 14 cM region in the terminal part of chromosome 13q that included the locus for COL4A1. The affected family members were heterozygous for a novel COL4A1 sequence variant c.4881C>G (p...
  6. Sibon I, Coupry I, Menegon P, Bouchet J, Gorry P, Burgelin I, et al. COL4A1 mutation in Axenfeld-Rieger anomaly with leukoencephalopathy and stroke. Ann Neurol. 2007;62:177-84 pubmed
    ..Familial genetic analyses led to the identification of a novel missense mutation in the COL4A1 gene, p.G720D, which cosegregates with the disease...
  7. Tarasov K, Sanna S, Scuteri A, Strait J, Orrù M, Parsa A, et al. COL4A1 is associated with arterial stiffness by genome-wide association scan. Circ Cardiovasc Genet. 2009;2:151-8 pubmed publisher
    ..Of the loci that were tested for association with PWV, the nonsynonymous SNP rs3742207 in the COL4A1 gene on chromosome 13 and SNP rs1495448 in the MAGI1 gene on chromosome 3 were successfully replicated (P=7...
  8. Jeanne M, Labelle Dumais C, Jorgensen J, Kauffman W, Mancini G, Favor J, et al. COL4A2 mutations impair COL4A1 and COL4A2 secretion and cause hemorrhagic stroke. Am J Hum Genet. 2012;90:91-101 pubmed publisher
    Collagen, type IV, alpha 1 (COL4A1) and alpha 2 (COL4A2) form heterotrimers and are abundant components of basement membranes, including those of the cerebral vasculature...
  9. Vermeulen R, Peeters Scholte C, Van Vugt J, Van Vught J, Barkhof F, Rizzu P, et al. Fetal origin of brain damage in 2 infants with a COL4A1 mutation: fetal and neonatal MRI. Neuropediatrics. 2011;42:1-3 pubmed publisher
    Mutations in the gene COL4A1, encoding collagen IV A1, are associated with familial porencephaly. Previously, COL4A1 mutation-associated antenatal hemorrhages have been suggested by early post-natal imaging...

More Information


  1. Nyberg P, Xie L, Sugimoto H, Colorado P, Sund M, Holthaus K, et al. Characterization of the anti-angiogenic properties of arresten, an alpha1beta1 integrin-dependent collagen-derived tumor suppressor. Exp Cell Res. 2008;314:3292-305 pubmed publisher
    ..Several collagen-derived fragments possess anti-angiogenic activity. Arresten is the 26-kDa non-collagenous domain of type IV collagen alpha1 chain...
  2. de Vries L, Koopman C, Groenendaal F, van Schooneveld M, Verheijen F, Verbeek E, et al. COL4A1 mutation in two preterm siblings with antenatal onset of parenchymal hemorrhage. Ann Neurol. 2009;65:12-8 pubmed publisher
    ..Mutation analysis of the COL4A1 gene was performed in the infants and in their mother...
  3. Sudhakar A, Nyberg P, Keshamouni V, Mannam A, Li J, Sugimoto H, et al. Human alpha1 type IV collagen NC1 domain exhibits distinct antiangiogenic activity mediated by alpha1beta1 integrin. J Clin Invest. 2005;115:2801-10 pubmed
    Human noncollagenous domain 1 of the alpha1 chain of type IV collagen [alpha1(IV)NC1], or arresten, is derived from the carboxy terminal of type IV collagen...
  4. Bilguvar K, DiLuna M, Bizzarro M, Bayri Y, Schneider K, Lifton R, et al. COL4A1 mutation in preterm intraventricular hemorrhage. J Pediatr. 2009;155:743-5 pubmed publisher
    Intraventricular hemorrhage is a common complication of preterm infants. Mutations in the type IV procollagen gene, COL4A1, are associated with cerebral small vessel disease with hemorrhage in adults and fetuses...
  5. Fischer G, Schmidt C, Opitz J, Cully Z, Kuhn K, Poschl E. Identification of a novel sequence element in the common promoter region of human collagen type IV genes, involved in the regulation of divergent transcription. Biochem J. 1993;292 ( Pt 3):687-95 pubmed
    ..The two genes COL4A1 and COL4A2 that code for the subunits are found closely linked on chromosome 13 in a head-to-head arrangement and ..
  6. Assadian S, El Assaad W, Wang X, Gannon P, Barrès V, Latour M, et al. p53 inhibits angiogenesis by inducing the production of Arresten. Cancer Res. 2012;72:1270-9 pubmed publisher
    ..Among those is Arresten, a collagen-derived antiangiogenic factor (CDAF) that is processed from ?1 collagen IV...
  7. Kuo D, Labelle Dumais C, Gould D. COL4A1 and COL4A2 mutations and disease: insights into pathogenic mechanisms and potential therapeutic targets. Hum Mol Genet. 2012;21:R97-110 pubmed
    Heterotrimers composed of collagen type IV alpha 1 (COL4A1) and alpha 2 (COL4A2) constitute one of the most abundant components of nearly all basement membranes...
  8. Than M, Henrich S, Huber R, Ries A, Mann K, Kuhn K, et al. The 1.9-A crystal structure of the noncollagenous (NC1) domain of human placenta collagen IV shows stabilization via a novel type of covalent Met-Lys cross-link. Proc Natl Acad Sci U S A. 2002;99:6607-12 pubmed
    ..This structure provides insights into NC1-related diseases such as Goodpasture and Alport syndromes. ..
  9. Meuwissen M, de Vries L, Verbeek H, Lequin M, Govaert P, Schot R, et al. Sporadic COL4A1 mutations with extensive prenatal porencephaly resembling hydranencephaly. Neurology. 2011;76:844-6 pubmed publisher
  10. Breedveld G, de Coo I, Lequin M, Arts W, Heutink P, Gould D, et al. Novel mutations in three families confirm a major role of COL4A1 in hereditary porencephaly. J Med Genet. 2006;43:490-5 pubmed
    ..b>COL4A1 is an essential component in basal membrane stability...
  11. Weng Y, Sonni A, Labelle Dumais C, de Leau M, Kauffman W, Jeanne M, et al. COL4A1 mutations in patients with sporadic late-onset intracerebral hemorrhage. Ann Neurol. 2012;71:470-7 pubmed publisher
    Mutations in the type IV collagen alpha 1 gene (COL4A1) cause dominantly inherited cerebrovascular disease. We seek to determine the extent to which COL4A1 mutations contribute to sporadic, nonfamilial, intracerebral hemorrhages (ICHs)...
  12. Shah S, Ellard S, Kneen R, Lim M, Osborne N, Rankin J, et al. Childhood presentation of COL4A1 mutations. Dev Med Child Neurol. 2012;54:569-74 pubmed publisher
    To describe the clinical and radiological features of four new families with a childhood presentation of COL4A1 mutation. We retrospectively reviewed the clinical presentation...
  13. Boosani C, Sudhakar A. Cloning, purification, and characterization of a non-collagenous anti-angiogenic protein domain from human alpha1 type IV collagen expressed in Sf9 cells. Protein Expr Purif. 2006;49:211-8 pubmed
    ..The purified protein showed its characteristic activities such as inhibiting cell proliferation, migration, and tube formation in endothelial cells. ..
  14. Coupry I, Sibon I, Mortemousque B, Rouanet F, Mine M, Goizet C. Ophthalmological features associated with COL4A1 mutations. Arch Ophthalmol. 2010;128:483-9 pubmed publisher
    To investigate the wide variability of ocular manifestations associated with mutations in the COL4A1 gene that encodes collagen IValpha1...
  15. Alamowitch S, Plaisier E, Favrole P, Prost C, Chen Z, Van Agtmael T, et al. Cerebrovascular disease related to COL4A1 mutations in HANAC syndrome. Neurology. 2009;73:1873-82 pubmed publisher
    b>COL4A1 mutations cause familial porencephaly, infantile hemiplegia, cerebral small vessel disease (CSVD), and hemorrhagic stroke...
  16. Yamada Y, Kato K, Oguri M, Fujimaki T, Yokoi K, Matsuo H, et al. Genetic risk for myocardial infarction determined by polymorphisms of candidate genes in a Japanese population. J Med Genet. 2008;45:216-21 pubmed
    ..Although several environmental factors influence the development of myocardial infarction (MI), genetic factors have been shown to contribute to individual susceptibility to this condition...
  17. van der Knaap M, Smit L, Barkhof F, Pijnenburg Y, Zweegman S, Niessen H, et al. Neonatal porencephaly and adult stroke related to mutations in collagen IV A1. Ann Neurol. 2006;59:504-11 pubmed
    ..Mutations in collagen IV A1, a major component of the vascular basement membrane, appear to be another risk factor. ..
  18. Vahedi K, Boukobza M, Massin P, Gould D, Tournier Lasserve E, Bousser M. Clinical and brain MRI follow-up study of a family with COL4A1 mutation. Neurology. 2007;69:1564-8 pubmed
    To better delineate the clinical spectrum and the natural history of COL4A1 mutations, a newly defined genetic cause of small vessel disease including the brain and retina...
  19. Gould D, Phalan F, van Mil S, Sundberg J, Vahedi K, Massin P, et al. Role of COL4A1 in small-vessel disease and hemorrhagic stroke. N Engl J Med. 2006;354:1489-96 pubmed
    ..In this report, we show that a mutation in the mouse Col4a1 gene, encoding procollagen type IV alpha1, predisposes both newborn and adult mice to intracerebral hemorrhage...
  20. Colorado P, Torre A, Kamphaus G, Maeshima Y, Hopfer H, Takahashi K, et al. Anti-angiogenic cues from vascular basement membrane collagen. Cancer Res. 2000;60:2520-6 pubmed
    ..basement membrane-associated protein, the 26-kDa NC1 domain of the alpha1 chain of type IV collagen, termed arresten. Arresten was isolated from human placenta and produced as a recombinant molecule in Escherichia coli and 293 ..
  21. Labelle Dumais C, Dilworth D, Harrington E, de Leau M, Lyons D, Kabaeva Z, et al. COL4A1 mutations cause ocular dysgenesis, neuronal localization defects, and myopathy in mice and Walker-Warburg syndrome in humans. PLoS Genet. 2011;7:e1002062 pubmed publisher
    ..we provide evidence that mutations in a gene coding for a major basement membrane protein, collagen IV alpha 1 (COL4A1), are a novel cause of MEB/WWS...
  22. Du B, Ma L, Huang M, Zhou H, Huang H, Shao P, et al. High glucose down-regulates miR-29a to increase collagen IV production in HK-2 cells. FEBS Lett. 2010;584:811-6 pubmed publisher
    ..We then showed that miR-29a negatively regulated collagen IV by directly targeting the 3'UTRs of col4a1 and col4a2...
  23. Shah S, Kumar Y, McLean B, Churchill A, Stoodley N, Rankin J, et al. A dominantly inherited mutation in collagen IV A1 (COL4A1) causing childhood onset stroke without porencephaly. Eur J Paediatr Neurol. 2010;14:182-7 pubmed publisher
    ..All of the family members had cataracts but did not have retinal arterial changes. Sequence analysis of COL4A1 revealed the heterozygous missense mutation c...
  24. Hinek A. Nature and the multiple functions of the 67-kD elastin-/laminin binding protein. Cell Adhes Commun. 1994;2:185-93 pubmed
    ..I will emphasize the role of the 67-kD protein in divergent cellular processes, such as cell-matrix attachment, matrix assembly, cellular chemotaxis, and active migration through the vascular walls. ..
  25. Mase Y, Ishibashi O, Ishikawa T, Takizawa T, Kiguchi K, Ohba T, et al. MiR-21 is enriched in the RNA-induced silencing complex and targets COL4A1 in human granulosa cell lines. Reprod Sci. 2012;19:1030-40 pubmed publisher
    ..We identified COL4A1 mRNA as a miR-21 target in the GC lines...
  26. Yoneda Y, Haginoya K, Kato M, Osaka H, Yokochi K, Arai H, et al. Phenotypic spectrum of COL4A1 mutations: porencephaly to schizencephaly. Ann Neurol. 2013;73:48-57 pubmed publisher
    Recently, COL4A1 mutations have been reported in porencephaly and other cerebral vascular diseases, often associated with ocular, renal, and muscular features...
  27. Xiao J, Sun X, Madhan B, Brodsky B, Baum J. NMR studies demonstrate a unique AAB composition and chain register for a heterotrimeric type IV collagen model peptide containing a natural interruption site. J Biol Chem. 2015;290:24201-9 pubmed publisher
    ..This study illustrates the potential role of interruptions in the structure, dynamics, and folding of natural collagen heterotrimers and forms a basis for understanding their biological role. ..
  28. Bager C, Willumsen N, Leeming D, Smith V, Karsdal M, Dornan D, et al. Collagen degradation products measured in serum can separate ovarian and breast cancer patients from healthy controls: A preliminary study. Cancer Biomark. 2015;15:783-8 pubmed publisher
    ..Four serum biomarkers reflecting altered MMP-mediated collagen turnover were able to differentiate ovarian and breast cancer patients from healthy controls. ..
  29. Gasparini S, Qualtieri A, Ferlazzo E, Cianci V, Patitucci A, Spadafora P, et al. Normal immunofluorescence pattern of skin basement membranes in a family with porencephaly due to COL4A1 G749S mutation. Neurol Sci. 2016;37:459-63 pubmed publisher
    b>COL4A1 mutations have been associated with cerebral small-vessel disease, including perinatal intracerebral hemorrhage with consequent porencephaly, microbleeds, and lacunar strokes...
  30. Smigiel R, Cabala M, Jakubiak A, Kodera H, Sasiadek M, Matsumoto N, et al. Novel COL4A1 mutation in an infant with severe dysmorphic syndrome with schizencephaly, periventricular calcifications, and cataract resembling congenital infection. Birth Defects Res A Clin Mol Teratol. 2016;106:304-7 pubmed publisher
    ..Chromosomal analysis and array-CGH showed no aberration. Target capture sequencing for COL4A1 and COL4A2 revealed a de novo COL4A1 mutation (c.2123G>T [p.Gly708Val])...
  31. Johnson G, Moore S. Human acetylcholinesterase binds to mouse laminin-1 and human collagen IV by an electrostatic mechanism at the peripheral anionic site. Neurosci Lett. 2003;337:37-40 pubmed
  32. Sasaki T, Göhring W, Pan T, Chu M, Timpl R. Binding of mouse and human fibulin-2 to extracellular matrix ligands. J Mol Biol. 1995;254:892-9 pubmed
    ..This indicated multifunctional binding properties for fibulin-2 and several alternative routes for its integration into basement membranes and other extracellular structures. ..
  33. Surlin P, Rauten A, Pirici D, Oprea B, Mogoanta L, Camen A. Collagen IV and MMP-9 expression in hypertrophic gingiva during orthodontic treatment. Rom J Morphol Embryol. 2012;53:161-5 pubmed
  34. Gale D, Oygar D, Lin F, Oygar P, Khan N, Connor T, et al. A novel COL4A1 frameshift mutation in familial kidney disease: the importance of the C-terminal NC1 domain of type IV collagen. Nephrol Dial Transplant. 2016;31:1908-1914 pubmed
    ..We identified a novel frameshift mutation, c.4611_4612insG:p.T1537fs, in exon 49 of COL4A1. This mutation predicts truncation of the protein with disruption of the C-terminal part of the NC1 domain...
  35. Cutting G, Kazazian H, Antonarakis S, Killen P, Yamada Y, Francomano C. Macrorestriction mapping of COL4A1 and COL4A2 collagen genes on human chromosome 13q34. Genomics. 1988;3:256-63 pubmed
    The genes for the alpha-1 and alpha-2 chains of type IV collagen (COL4A1 and COL4A2) map to the same chromosomal band (13q34) and have a high degree of nucleotide homology...
  36. Chen M, Marinkovich M, Veis A, Cai X, Rao C, O Toole E, et al. Interactions of the amino-terminal noncollagenous (NC1) domain of type VII collagen with extracellular matrix components. A potential role in epidermal-dermal adherence in human skin. J Biol Chem. 1997;272:14516-22 pubmed
    ..The multiple interactions of NC1 with other extracellular matrix components may support epidermal-dermal adhesion. ..
  37. Sundarraj N, Willson J. Monoclonal antibody to human basement membrane collagen type IV. Immunology. 1982;47:133-40 pubmed
    ..Based on the observations of other investigators, these peptides are probably derived by proteolysis of the larger polypeptides, since the purification steps in involve extensive pepsin digestion. ..
  38. Kusuma S, Zhao S, Gerecht S. The extracellular matrix is a novel attribute of endothelial progenitors and of hypoxic mature endothelial cells. FASEB J. 2012;26:4925-36 pubmed publisher
    ..Collectively, we suggest that ECM deposition and assembly by ECs is dependent on maturation stage and oxygen supply and that these findings can be harnessed to advance engineered vascular therapeutics. ..
  39. Ruigrok Y, Rinkel G, Wijmenga C, Kasuya H, Tajima A, Takahashi T, et al. Association analysis of genes involved in the maintenance of the integrity of the extracellular matrix with intracranial aneurysms in a Japanese cohort. Cerebrovasc Dis. 2009;28:131-4 pubmed publisher
    An association between versican (CSPG2), perlecan (HSPG2), fibrillin 2 (FBN2) and collagen 4A1 (COL4A1) gene variants and intracranial aneurysms (IA) has been reported in 2 studies analyzing Dutch IA patients...
  40. Grosfeld A, Stolze I, Cockman M, Pugh C, Edelmann M, Kessler B, et al. Interaction of hydroxylated collagen IV with the von hippel-lindau tumor suppressor. J Biol Chem. 2007;282:13264-9 pubmed
    ..The interaction was suppressed by a wide range of tumor-associated mutations, including those that do not disturb the regulation of HIF, supporting a role in HIF-independent tumor suppressor functions. ..
  41. Babel W, Glanville R. Structure of human-basement-membrane (type IV) collagen. Complete amino-acid sequence of a 914-residue-long pepsin fragment from the alpha 1(IV) chain. Eur J Biochem. 1984;143:545-56 pubmed
    ..The unique sequence characteristics of type IV collagen are discussed in relation to its macromolecular structure and to the interstitial collagens...
  42. Aumailley M, Wiedemann H, Mann K, Timpl R. Binding of nidogen and the laminin-nidogen complex to basement membrane collagen type IV. Eur J Biochem. 1989;184:241-8 pubmed
    ..A more diverse collagen-IV-binding pattern was observed for the laminin nidogen complex, whereby interactions may involve both nidogen and short-arm structures of laminin. ..
  43. Liao J, Su X, Chen P, Wang X, Xu L, Li X, et al. Meta-analysis of genome-wide association studies in multiethnic Asians identifies two loci for age-related nuclear cataract. Hum Mol Genet. 2014;23:6119-28 pubmed publisher
    ..The results offer additional insights into the pathogenesis of nuclear cataract in Asians. ..
  44. Glanville R, Qian R, Siebold B, Risteli J, Kuhn K. Amino acid sequence of the N-terminal aggregation and cross-linking region (7S domain) of the alpha 1 (IV) chain of human basement membrane collagen. Eur J Biochem. 1985;152:213-9 pubmed
    ..The mode of aggregation of the 7S domain, the formation of intermolecular cross-links as well as the relatively high stability of this region against proteolytic attack are discussed in the light of the elucidated amino acid sequence. ..
  45. Li X, Wu Z, Ni J, Liu Y, Meng J, Yu W, et al. Cathepsin B Regulates Collagen Expression by Fibroblasts via Prolonging TLR2/NF-?B Activation. Oxid Med Cell Longev. 2016;2016:7894247 pubmed publisher
    ..CatB-specific inhibitors may therefore improve chronic inflammation-delayed tissue repair. ..
  46. Januszyk M, Kwon S, Wong V, Padmanabhan J, Maan Z, Whittam A, et al. The Role of Focal Adhesion Kinase in Keratinocyte Fibrogenic Gene Expression. Int J Mol Sci. 2017;18: pubmed publisher
    ..of genes integral to mechanotransduction and extracellular matrix production, including Igtbl, Mmpla, and Col4a1. Transcriptional activities upon FAK deletion were not identical across all single keratinocytes, resulting in ..
  47. Boyd C, Weliky K, Toth Fejel S, Deak S, Christiano A, Mackenzie J, et al. The single copy gene coding for human alpha 1 (IV) procollagen is located at the terminal end of the long arm of chromosome 13. Hum Genet. 1986;74:121-5 pubmed
    ..In addition, Southern and slot blot hybridization analysis clearly show that these genomic sequences are present only once per haploid genome. ..
  48. Pihlajaniemi T, Tryggvason K, Myers J, Kurkinen M, Lebo R, Cheung M, et al. cDNA clones coding for the pro-alpha1(IV) chain of human type IV procollagen reveal an unusual homology of amino acid sequences in two halves of the carboxyl-terminal domain. J Biol Chem. 1985;260:7681-7 pubmed
    ..In further studies, sorted human chromosomes were used to locate the gene for the pro-alpha1(IV) chain on chromosome 13. ..
  49. Nyström H, Tavelin B, Björklund M, Naredi P, Sund M. Improved tumour marker sensitivity in detecting colorectal liver metastases by combined type IV collagen and CEA measurement. Tumour Biol. 2015;36:9839-47 pubmed publisher
    ..001). COLIV is a promising tumour marker for CLM and can possibly be used to detect postoperative CLM recurrence. The combination of COLIV and CEA is superior to either marker alone in detecting CLM. ..
  50. Chen S, Lin J, Lin H, Shan Y, Cheng Y, Yang B. Dependence of fibroblast infiltration in tumor stroma on type IV collagen-initiated integrin signal through induction of platelet-derived growth factor. Biochim Biophys Acta. 2015;1853:929-39 pubmed publisher
    ..The integrin signal pathway was activated by collagen engagement through Src and ERK, leading to enhanced PDGF-A production, which serves as a key regulator of fibroblast recruitment. ..
  51. Vahedi K, Alamowitch S. Clinical spectrum of type IV collagen (COL4A1) mutations: a novel genetic multisystem disease. Curr Opin Neurol. 2011;24:63-8 pubmed publisher
    This review dsecribes the clinical spectrum of a newly identified disorder related to COL4A1 gene mutations...
  52. Cortes Reynosa P, Robledo T, Macias Silva M, Wu S, Salazar E. Src kinase regulates metalloproteinase-9 secretion induced by type IV collagen in MCF-7 human breast cancer cells. Matrix Biol. 2008;27:220-31 pubmed
    ..In contrast, MMP-9 expression was not up-regulated by treatment with Col-IV. These results demonstrate that Col-IV regulates the secretion of MMP-9 via a Src and FAK dependent pathway in MCF-7 cells. ..
  53. Soininen R, Haka Risku T, Prockop D, Tryggvason K. Complete primary structure of the alpha 1-chain of human basement membrane (type IV) collagen. FEBS Lett. 1987;225:188-94 pubmed
    ..The alpha 1(IV)-chain contains four cysteine residues in the triple-helical domain, four cysteines in the 15-residue long noncollagenous sequence at the amino-terminus and 12 cysteines in the carboxy-terminal NC-domain. ..