{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T16:15:27Z","timestamp":1791389727994,"version":"4.3.3"},"reference-count":56,"publisher":"The Endocrine Society","issue":"8","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2009,8,1]]},"abstract":"<jats:p>GNAS gives rise to multiple imprinted gene products, including the \u03b1-subunit of the stimulatory G protein (Gs\u03b1) and its variant XL\u03b1s. Based on genomic sequence, the translation of XL\u03b1s begins from the middle of a long open reading frame, suggesting the existence of an N-terminally extended variant termed extralarge XL\u03b1s (XXL\u03b1s). Although XXL\u03b1s, like Gs\u03b1 and XL\u03b1s, would be affected by most disease-causing GNAS mutations, its authenticity and biological significance remained unknown. Here we identified a mouse cDNA clone that comprises the entire open reading frame encoding XXL\u03b1s. Whereas XXL\u03b1s mRNA was readily detected in mouse heart by RT-PCR, it appeared virtually absent in insulinoma-derived INS-1 cells. By Northern blots and RT-PCR, XXL\u03b1s mRNA was detected primarily in the mouse brain, cerebellum, and spleen. Immunohistochemistry using a specific anti-XXL\u03b1s antibody demonstrated XXL\u03b1s protein in multiple brain areas, including dorsal hippocampus and cortex. In transfected cells, full-length human XXL\u03b1s was localized to the plasma membrane and mediated isoproterenol- and cholera toxin-stimulated cAMP accumulation. XXL\u03b1s-R844H, which bears a mutation analogous to that in the constitutively active Gs\u03b1 mutant Gs\u03b1-R201H (gsp oncogene), displayed elevated basal signaling. However, unlike Gs\u03b1-R201H, which mostly remains in the cytoplasm, both XXL\u03b1s-R844H and a constitutively active XL\u03b1s mutant localized to the plasma membrane. Hence, XXL\u03b1s is a distinct GNAS product and can mimic Gs\u03b1, but the constitutively active XXL\u03b1s and Gs\u03b1 mutants differ from each other regarding subcellular targeting. Our findings suggest that XXL\u03b1s deficiency or hyperactivity may contribute to the pathogenesis of diseases caused by GNAS mutations.<\/jats:p>","DOI":"10.1210\/en.2009-0318","type":"journal-article","created":{"date-parts":[[2009,5,8]],"date-time":"2009-05-08T01:30:34Z","timestamp":1241746234000},"page":"3567-3575","source":"Crossref","is-referenced-by-count":30,"title":["Extralarge XL\u03b1s (XXL\u03b1s), a Variant of Stimulatory G Protein \u03b1-Subunit (Gs\u03b1), Is a Distinct, Membrane-Anchored GNAS Product that Can Mimic Gs\u03b1"],"prefix":"10.1210","volume":"150","author":[{"given":"Cumhur","family":"Aydin","sequence":"first","affiliation":[{"name":"Department of Endocrine Unit (C.A., M.J.M., H.A.W.T., M.B.), Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts 02114"},{"name":"Department of Endodontics (C.A.), Center for Dental Sciences, Gu\u0308lhane School of Medicine, 06018 Ankara, Turkey"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nurgul","family":"Aytan","sequence":"first","affiliation":[{"name":"Departments of Neurology (N.A., N.W.K., A.D.), Boston University School of Medicine, Boston, Massachusetts 02115"},{"name":"Research and Development Service\/Geriatric Research Education and Clinical Center (N.A., N.W.K., A.D.), Veterans Affairs Boston Healthcare System, Boston, Massachusetts 02130"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mathew J.","family":"Mahon","sequence":"first","affiliation":[{"name":"Department of Endocrine Unit (C.A., M.J.M., H.A.W.T., M.B.), Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts 02114"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hesham A. W.","family":"Tawfeek","sequence":"first","affiliation":[{"name":"Department of Endocrine Unit (C.A., M.J.M., H.A.W.T., M.B.), Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts 02114"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Neil W.","family":"Kowall","sequence":"first","affiliation":[{"name":"Departments of Neurology (N.A., N.W.K., A.D.), Boston University School of Medicine, Boston, Massachusetts 02115"},{"name":"Departments of Pathology (N.W.K.), Boston University School of Medicine, Boston, Massachusetts 02115"},{"name":"Research and Development Service\/Geriatric Research Education and Clinical Center (N.A., N.W.K., A.D.), Veterans Affairs Boston Healthcare System, Boston, Massachusetts 02130"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alpaslan","family":"Dedeoglu","sequence":"first","affiliation":[{"name":"Departments of Neurology (N.A., N.W.K., A.D.), Boston University School of Medicine, Boston, Massachusetts 02115"},{"name":"Research and Development Service\/Geriatric Research Education and Clinical Center (N.A., N.W.K., A.D.), Veterans Affairs Boston Healthcare System, Boston, Massachusetts 02130"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Murat","family":"Bastepe","sequence":"first","affiliation":[{"name":"Department of Endocrine Unit (C.A., M.J.M., H.A.W.T., M.B.), Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts 02114"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"80","published-online":{"date-parts":[[2009,5,7]]},"reference":[{"key":"2020071612264998300_R1","doi-asserted-by":"crossref","first-page":"5459","DOI":"10.1210\/en.2004-0865","article-title":"Minireview: GNAS: normal and abnormal 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locus.","volume":"10","author":"Bastepe","year":"2001","journal-title":"Hum Mol Genet"},{"key":"2020071612264998300_R27","doi-asserted-by":"crossref","first-page":"2370","DOI":"10.1210\/jc.2006-2287","article-title":"Epigenetic defects of GNAS in patients with pseudohypoparathyroidism and mild features of Albright\u2019s hereditary osteodystrophy.","volume":"92","author":"Nanclares","year":"2007","journal-title":"J Clin Endocrinol Metab"},{"key":"2020071612264998300_R28","first-page":"1372","article-title":"Processing of gene expression data generated by quantitative real-time RT-PCR.","volume":"32","author":"Muller","year":"2002","journal-title":"Biotechniques"},{"key":"2020071612264998300_R29","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.brainres.2008.01.095","article-title":"Ibuprofen reduces A\u03b2, hyperphosphorylated \u03c4 and memory deficits in Alzheimer mice.","volume":"1207","author":"McKee","year":"2008","journal-title":"Brain 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pathways.","volume":"16","author":"Kovalovsky","year":"2002","journal-title":"Mol Endocrinol"},{"key":"2020071612264998300_R36","doi-asserted-by":"crossref","first-page":"1297","DOI":"10.1083\/jcb.119.5.1297","article-title":"Activation of the \u03b1 subunit of Gs in intact cells alters its abundance, rate of degradation, and membrane avidity.","volume":"119","author":"Levis","year":"1992","journal-title":"J Cell Biol"},{"key":"2020071612264998300_R37","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1016\/0092-8674(94)90445-6","article-title":"Activation and depalmitoylation of Gs\u03b1.","volume":"77","author":"Wedegaertner","year":"1994","journal-title":"Cell"},{"key":"2020071612264998300_R38","doi-asserted-by":"crossref","first-page":"1688","DOI":"10.1056\/NEJM199112123252403","article-title":"Activating mutations of the stimulatory G protein in the McCune-Albright syndrome.","volume":"325","author":"Weinstein","year":"1991","journal-title":"N Engl J 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