{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T14:09:58Z","timestamp":1791209398738,"version":"4.1.0"},"reference-count":52,"publisher":"Elsevier BV","issue":"41","license":[{"start":{"date-parts":[[2012,10,1]],"date-time":"2012-10-01T00:00:00Z","timestamp":1349049600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2020,9,16]],"date-time":"2020-09-16T00:00:00Z","timestamp":1600214400000},"content-version":"vor","delay-in-days":2907,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Journal of Biological Chemistry"],"published-print":{"date-parts":[[2012,10]]},"DOI":"10.1074\/jbc.m112.381970","type":"journal-article","created":{"date-parts":[[2012,8,11]],"date-time":"2012-08-11T22:01:45Z","timestamp":1344722505000},"page":"34167-34178","source":"Crossref","is-referenced-by-count":121,"title":["Precise Regulation of Porcupine Activity Is Required for Physiological Wnt Signaling"],"prefix":"10.1016","volume":"287","author":[{"given":"Kyle D.","family":"Proffitt","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David M.","family":"Virshup","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1074\/jbc.M112.381970_bib1","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.devcel.2009.06.016","article-title":"Wnt\/beta-catenin signaling. Components, mechanisms, and diseases","volume":"17","author":"MacDonald","year":"2009","journal-title":"Dev. Cell"},{"key":"10.1074\/jbc.M112.381970_bib2","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1359\/jbmr.081235","article-title":"Where Wnts went. The exploding field of Lrp5 and Lrp6 signaling in bone","volume":"24","author":"Williams","year":"2009","journal-title":"J. Bone Miner. Res."},{"key":"10.1074\/jbc.M112.381970_bib3","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.gde.2006.12.007","article-title":"The many ways of Wnt in cancer","volume":"17","author":"Polakis","year":"2007","journal-title":"Curr. Opin. Genet. Dev."},{"key":"10.1074\/jbc.M112.381970_bib4","doi-asserted-by":"crossref","first-page":"2676","DOI":"10.2353\/ajpath.2009.080945","article-title":"Activation of the Wnt pathway plays a pathogenic role in diabetic retinopathy in humans and animal models","volume":"175","author":"Chen","year":"2009","journal-title":"Am. J. Pathol."},{"key":"10.1074\/jbc.M112.381970_bib5","doi-asserted-by":"crossref","first-page":"562","DOI":"10.1097\/BOR.0b013e32834b3309","article-title":"\u03b2-Catenin signaling. A novel mediator of fibrosis and potential therapeutic target","volume":"23","author":"Lam","year":"2011","journal-title":"Curr. Opin. Rheumatol."},{"key":"10.1074\/jbc.M112.381970_bib6","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1016\/S0092-8674(00)81991-1","article-title":"Direct and long range action of a wingless morphogen gradient","volume":"87","author":"Zecca","year":"1996","journal-title":"Cell"},{"key":"10.1074\/jbc.M112.381970_bib7","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1016\/j.devcel.2011.01.001","article-title":"Wnt signaling gradients establish planar cell polarity by inducing Vangl2 phosphorylation through Ror2","volume":"20","author":"Gao","year":"2011","journal-title":"Dev. Cell"},{"key":"10.1074\/jbc.M112.381970_bib8","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1038\/embor.2008.167","article-title":"Regulation of Wnt protein secretion and its role in gradient formation","volume":"9","author":"Bartscherer","year":"2008","journal-title":"EMBO Rep."},{"key":"10.1074\/jbc.M112.381970_bib9","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1111\/j.1748-1716.2011.02287.x","article-title":"Dissecting the Wnt secretion pathway. Key questions on the modification and intracellular trafficking of Wnt proteins","volume":"204","author":"Harterink","year":"2012","journal-title":"Acta. Physiol."},{"key":"10.1074\/jbc.M112.381970_bib10","doi-asserted-by":"crossref","first-page":"1144","DOI":"10.1038\/embor.2011.165","article-title":"A genome-wide RNA interference screen uncovers two p24 proteins as regulators of Wingless secretion","volume":"12","author":"Port","year":"2011","journal-title":"EMBO Rep."},{"key":"10.1074\/jbc.M112.381970_bib11","doi-asserted-by":"crossref","first-page":"1265","DOI":"10.1038\/embor.2011.212","article-title":"p24 proteins are required for secretion of Wnt ligands","volume":"12","author":"Buechling","year":"2011","journal-title":"EMBO Rep."},{"key":"10.1074\/jbc.M112.381970_bib12","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1016\/j.cell.2006.02.049","article-title":"Wntless, a conserved membrane protein dedicated to the secretion of Wnt proteins from signaling cells","volume":"125","author":"B\u00e4nziger","year":"2006","journal-title":"Cell"},{"key":"10.1074\/jbc.M112.381970_bib13","doi-asserted-by":"crossref","first-page":"523","DOI":"10.1016\/j.cell.2006.04.009","article-title":"Secretion of Wnt ligands requires Evi, a conserved transmembrane protein","volume":"125","author":"Bartscherer","year":"2006","journal-title":"Cell"},{"key":"10.1074\/jbc.M112.381970_bib14","doi-asserted-by":"crossref","first-page":"4901","DOI":"10.1242\/dev.02674","article-title":"Sprinter. A novel transmembrane protein required for Wg secretion and signaling","volume":"133","author":"Goodman","year":"2006","journal-title":"Development"},{"key":"10.1074\/jbc.M112.381970_bib15","doi-asserted-by":"crossref","first-page":"914","DOI":"10.1038\/ncb2281","article-title":"A SNX3-dependent retromer pathway mediates retrograde transport of the Wnt sorting receptor Wntless and is required for Wnt secretion","volume":"13","author":"Harterink","year":"2011","journal-title":"Nat. Cell Biol."},{"key":"10.1074\/jbc.M112.381970_bib16","doi-asserted-by":"crossref","first-page":"3116","DOI":"10.1101\/gad.10.24.3116","article-title":"The segment polarity gene porcupine encodes a putative multitransmembrane protein involved in Wingless processing","volume":"10","author":"Kadowaki","year":"1996","journal-title":"Genes Dev."},{"key":"10.1074\/jbc.M112.381970_bib17","doi-asserted-by":"crossref","first-page":"33220","DOI":"10.1074\/jbc.M403407200","article-title":"Drosophila wnt-1 undergoes a hydrophobic modification and is targeted to lipid rafts, a process that requires porcupine","volume":"279","author":"Zhai","year":"2004","journal-title":"J. Biol. Chem."},{"key":"10.1074\/jbc.M112.381970_bib18","doi-asserted-by":"crossref","first-page":"791","DOI":"10.1016\/j.devcel.2006.10.003","article-title":"Monounsaturated fatty acid modification of Wnt protein. Its role in Wnt secretion","volume":"11","author":"Takada","year":"2006","journal-title":"Dev Cell"},{"key":"10.1074\/jbc.M112.381970_bib19","doi-asserted-by":"crossref","first-page":"3357","DOI":"10.1242\/jcs.072132","article-title":"WLS-dependent secretion of WNT3A requires Ser-209 acylation and vacuolar acidification","volume":"123","author":"Coombs","year":"2010","journal-title":"J. Cell Sci."},{"key":"10.1074\/jbc.M112.381970_bib20","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1016\/j.ydbio.2011.11.003","article-title":"Porcupine-mediated lipidation is required for Wnt recognition by Wls","volume":"361","author":"Herr","year":"2012","journal-title":"Dev. Biol."},{"key":"10.1074\/jbc.M112.381970_bib21","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1126\/science.1222879","article-title":"Structural basis of Wnt recognition by Frizzled","volume":"337","author":"Janda","year":"2012","journal-title":"Science"},{"key":"10.1074\/jbc.M112.381970_bib22","doi-asserted-by":"crossref","first-page":"448","DOI":"10.1038\/nature01611","article-title":"Wnt proteins are lipid-modified and can act as stem cell growth factors","volume":"423","author":"Willert","year":"2003","journal-title":"Nature"},{"key":"10.1074\/jbc.M112.381970_bib23","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1042\/BJ20061476","article-title":"Post-translational palmitoylation and glycosylation of Wnt-5a are necessary for its signaling","volume":"402","author":"Kurayoshi","year":"2007","journal-title":"Biochem. J."},{"key":"10.1074\/jbc.M112.381970_bib24","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/S0968-0004(99)01539-X","article-title":"A superfamily of membrane-bound O-acyltransferases with implications for wnt signaling","volume":"25","author":"Hofmann","year":"2000","journal-title":"Trends Biochem. Sci."},{"key":"10.1074\/jbc.M112.381970_bib25","first-page":"2224","article-title":"Loss of Porcupine impairs convergent extension during gastrulation in zebrafish","volume":"125","author":"Chen","year":"2012","journal-title":"J. Cell Sci."},{"key":"10.1074\/jbc.M112.381970_bib26","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1016\/j.ydbio.2011.04.029","article-title":"Porcupine homolog is required for canonical Wnt signaling and gastrulation in mouse embryos","volume":"355","author":"Biechele","year":"2011","journal-title":"Dev. Biol."},{"key":"10.1074\/jbc.M112.381970_bib27","doi-asserted-by":"crossref","first-page":"12752","DOI":"10.1073\/pnas.1006437108","article-title":"Deletion of mouse Porcn blocks Wnt ligand secretion and reveals an ectodermal etiology of human focal dermal hypoplasia\/Goltz syndrome","volume":"108","author":"Barrott","year":"2011","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"10.1074\/jbc.M112.381970_bib28","doi-asserted-by":"crossref","first-page":"e32331","DOI":"10.1371\/journal.pone.0032331","article-title":"Deletion of Porcn in mice leads to multiple developmental defects and models human focal dermal hypoplasia (Goltz syndrome)","volume":"7","author":"Liu","year":"2012","journal-title":"PLoS ONE"},{"key":"10.1074\/jbc.M112.381970_bib29","doi-asserted-by":"crossref","first-page":"4300","DOI":"10.1046\/j.1432-1033.2000.01478.x","article-title":"The evolutionarily conserved porcupine gene family is involved in the processing of the Wnt family","volume":"267","author":"Tanaka","year":"2000","journal-title":"Eur. J. Biochem."},{"key":"10.1074\/jbc.M112.381970_bib30","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1016\/S0378-1119(02)00467-5","article-title":"Molecular cloning and initial characterization of the MG61\/PORC gene, the human homologue of the Drosophila segment polarity gene Porcupine","volume":"288","author":"Caricasole","year":"2002","journal-title":"Gene"},{"key":"10.1074\/jbc.M112.381970_bib31","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1038\/nchembio.137","article-title":"Small molecule-mediated disruption of Wnt-dependent signaling in tissue regeneration and cancer","volume":"5","author":"Chen","year":"2009","journal-title":"Nat. Chem. Biol."},{"key":"10.1074\/jbc.M112.381970_bib32","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1038\/nature07935","article-title":"Single Lgr5 stem cells build crypt-villus structures in vitro without a mesenchymal niche","volume":"459","author":"Sato","year":"2009","journal-title":"Nature"},{"key":"10.1074\/jbc.M112.381970_bib33","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1038\/ng2052","article-title":"Deficiency of PORCN, a regulator of Wnt signaling, is associated with focal dermal hypoplasia","volume":"39","author":"Grzeschik","year":"2007","journal-title":"Nat. Genet."},{"key":"10.1074\/jbc.M112.381970_bib34","doi-asserted-by":"crossref","first-page":"836","DOI":"10.1038\/ng2057","article-title":"Mutations in X-linked PORCN, a putative regulator of Wnt signaling, cause focal dermal hypoplasia","volume":"39","author":"Wang","year":"2007","journal-title":"Nat. Genet."},{"key":"10.1074\/jbc.M112.381970_bib35","doi-asserted-by":"crossref","first-page":"723","DOI":"10.1002\/humu.21505","article-title":"Mutation update for the PORCN gene","volume":"32","author":"Lombardi","year":"2011","journal-title":"Hum. Mutat."},{"key":"10.1074\/jbc.M112.381970_bib36","doi-asserted-by":"crossref","first-page":"e34532","DOI":"10.1371\/journal.pone.0034532","article-title":"PORCN moonlights in a Wnt-independent pathway that regulates cancer cell proliferation","volume":"7","author":"Covey","year":"2012","journal-title":"PLoS ONE"},{"key":"10.1074\/jbc.M112.381970_bib37","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1016\/S0960-9822(03)00240-9","article-title":"Zebrafish prickle, a modulator of noncanonical Wnt\/Fz signaling, regulates gastrulation movements","volume":"13","author":"Veeman","year":"2003","journal-title":"Curr. Biol."},{"key":"10.1074\/jbc.M112.381970_bib38","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.diff.2012.06.004","article-title":"A uniform human Wnt expression library reveals a shared secretory pathway and unique signaling activities","volume":"84","author":"Najdi","year":"2012","journal-title":"Differentiation"},{"key":"10.1074\/jbc.M112.381970_bib39","doi-asserted-by":"crossref","first-page":"3350","DOI":"10.1158\/0008-5472.CAN-07-3220","article-title":"Frzb, a secreted Wnt antagonist, decreases growth and invasiveness of fibrosarcoma cells associated with inhibition of Met signaling","volume":"68","author":"Guo","year":"2008","journal-title":"Cancer Res."},{"key":"10.1074\/jbc.M112.381970_bib40","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1016\/j.ccr.2011.03.010","article-title":"High frequency canonical Wnt activation in multiple sarcoma subtypes drives proliferation through a TCF\/\u03b2-catenin target gene, CDC25A","volume":"19","author":"Vijayakumar","year":"2011","journal-title":"Cancer Cell"},{"key":"10.1074\/jbc.M112.381970_bib41","doi-asserted-by":"crossref","first-page":"3483","DOI":"10.1038\/sj.onc.1211006","article-title":"Suppression of PPN\/MG61 attenuates Wnt\/\u03b2-catenin signaling pathway and induces apoptosis in human lung cancer","volume":"27","author":"Chen","year":"2008","journal-title":"Oncogene"},{"key":"10.1074\/jbc.M112.381970_bib42","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1016\/j.cell.2008.01.017","article-title":"Identification of the acyltransferase that octanoylates ghrelin, an appetite-stimulating peptide hormone","volume":"132","author":"Yang","year":"2008","journal-title":"Cell"},{"key":"10.1074\/jbc.M112.381970_bib43","doi-asserted-by":"crossref","first-page":"656","DOI":"10.1016\/j.stem.2010.11.016","article-title":"Tissue-resident adult stem cell populations of rapidly self-renewing organs","volume":"7","author":"Barker","year":"2010","journal-title":"Cell Stem Cell"},{"key":"10.1074\/jbc.M112.381970_bib44","doi-asserted-by":"crossref","first-page":"E618","DOI":"10.1002\/humu.20992","article-title":"PORCN mutations in focal dermal hypoplasia. Coping with lethality","volume":"30","author":"Bornholdt","year":"2009","journal-title":"Hum. Mutat."},{"key":"10.1074\/jbc.M112.381970_bib45","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1089\/gtmb.2010.0089","article-title":"PORCN mutations and variants identified in patients with focal dermal hypoplasia through diagnostic gene sequencing","volume":"14","author":"Fernandes","year":"2010","journal-title":"Genet. Test Mol. Biomarkers"},{"key":"10.1074\/jbc.M112.381970_bib46","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1111\/j.1399-0004.2009.01248.x","article-title":"Novel PORCN mutations in focal dermal hypoplasia","volume":"76","author":"Froyen","year":"2009","journal-title":"Clin. Genet."},{"key":"10.1074\/jbc.M112.381970_bib47","doi-asserted-by":"crossref","first-page":"1207","DOI":"10.1038\/ejhg.2009.40","article-title":"Goltz-Gorlin (focal dermal hypoplasia) and the microphthalmia with linear skin defects (MLS) syndrome. No evidence of genetic overlap","volume":"17","author":"Harmsen","year":"2009","journal-title":"Eur. J. Hum. Genet."},{"key":"10.1074\/jbc.M112.381970_bib48","doi-asserted-by":"crossref","first-page":"3339","DOI":"10.1242\/dev.02881","article-title":"Porcupine-mediated lipid modification regulates the activity and distribution of Wnt proteins in the chick neural tube","volume":"134","author":"Galli","year":"2007","journal-title":"Development"},{"key":"10.1074\/jbc.M112.381970_bib49","doi-asserted-by":"crossref","first-page":"e26636","DOI":"10.1371\/journal.pone.0026636","article-title":"Differential palmit(e)oylation of Wnt1 on Cys-93 and Ser-224 residues has overlapping and distinct consequences","volume":"6","author":"Galli","year":"2011","journal-title":"PLoS ONE"},{"key":"10.1074\/jbc.M112.381970_bib50","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1016\/j.cellsig.2011.01.007","article-title":"Fatty acid modification of Wnt1 and Wnt3a at serine is prerequisite for lipidation at cysteine and is essential for Wnt signaling","volume":"23","author":"Doubravska","year":"2011","journal-title":"Cell. Signal."},{"key":"10.1074\/jbc.M112.381970_bib51","doi-asserted-by":"crossref","first-page":"955","DOI":"10.1038\/nature07191","article-title":"The Trichoplax genome and the nature of placozoans","volume":"454","author":"Srivastava","year":"2008","journal-title":"Nature"},{"key":"10.1074\/jbc.M112.381970_bib52","doi-asserted-by":"crossref","first-page":"2830","DOI":"10.1073\/pnas.0712245105","article-title":"Discovery of a lysophospholipid acyltransferase family essential for membrane asymmetry and diversity","volume":"105","author":"Hishikawa","year":"2008","journal-title":"Proc. Natl. Acad. Sci. U.S.A."}],"container-title":["Journal of Biological Chemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925820627521?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925820627521?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1074\/jbc.M112.381970","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,12,18]],"date-time":"2021-12-18T10:10:15Z","timestamp":1639822215000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0021925820627521"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2012,10]]},"references-count":52,"journal-issue":{"issue":"41","published-print":{"date-parts":[[2012,10]]}},"alternative-id":["S0021925820627521"],"URL":"https:\/\/doi.org\/10.1074\/jbc.m112.381970","relation":{},"ISSN":["0021-9258"],"issn-type":[{"value":"0021-9258","type":"print"}],"subject":[],"published":{"date-parts":[[2012,10]]}}}