{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T03:54:13Z","timestamp":1792727653686,"version":"4.5.0"},"reference-count":44,"publisher":"American Society for Microbiology","issue":"4","license":[{"start":{"date-parts":[[2010,2,15]],"date-time":"2010-02-15T00:00:00Z","timestamp":1266192000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/journals.asm.org\/non-commercial-tdm-license"}],"content-domain":{"domain":["journals.asm.org"],"crossmark-restriction":true},"short-container-title":["J Virol"],"published-print":{"date-parts":[[2010,2,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n          <jats:p>The E5 protein of human papillomavirus type 16 is a small, hydrophobic protein that localizes predominantly to membranes of the endoplasmic reticulum (ER). To define the orientation of E5 in these membranes, we employed a differential, detergent permeabilization technique that makes use of the ability of low concentrations of digitonin to selectively permeabilize the plasma membrane and saponin to permeabilize all cellular membranes. We then generated a biologically active E5 protein that was epitope tagged at both its N and C termini and determined the accessibility of these termini to antibodies in the presence and absence of detergents. In both COS cells and human ectocervical cells, the C terminus of E5 was exposed to the cytoplasm, whereas the N terminus was restricted to the lumen of the ER. Finally, the deletion of the E5 third transmembrane domain (and terminal hydrophilic amino acids) resulted in a protein with its C terminus in the ER lumen. Taken together, these topology findings are compatible with a model of E5 being a 3-pass transmembrane protein and with studies demonstrating its C terminus interacting with cytoplasmic proteins.<\/jats:p>","DOI":"10.1128\/jvi.01968-09","type":"journal-article","created":{"date-parts":[[2009,12,3]],"date-time":"2009-12-03T03:25:20Z","timestamp":1259810720000},"page":"1696-1703","update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":45,"title":["Membrane Orientation of the Human Papillomavirus Type 16 E5 Oncoprotein"],"prefix":"10.1128","volume":"84","author":[{"given":"Ewa","family":"Krawczyk","sequence":"first","affiliation":[{"name":"Department of Pathology, Georgetown University Medical School, 3900 Reservoir Road NW, Washington, DC 20057"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Frank A.","family":"Suprynowicz","sequence":"additional","affiliation":[{"name":"Department of Pathology, Georgetown University Medical School, 3900 Reservoir Road NW, Washington, DC 20057"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sawali R.","family":"Sudarshan","sequence":"additional","affiliation":[{"name":"Department of Pathology, Georgetown University Medical School, 3900 Reservoir Road NW, Washington, DC 20057"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard","family":"Schlegel","sequence":"additional","affiliation":[{"name":"Department of Pathology, Georgetown University Medical School, 3900 Reservoir Road NW, Washington, DC 20057"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"235","reference":[{"key":"e_1_3_2_2_2","doi-asserted-by":"publisher","DOI":"10.1002\/ijc.22089"},{"key":"e_1_3_2_3_2","doi-asserted-by":"publisher","DOI":"10.4161\/cc.4.11.2152"},{"key":"e_1_3_2_4_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0002-9440(10)62552-7"},{"key":"e_1_3_2_5_2","doi-asserted-by":"publisher","DOI":"10.1016\/0968-0004(94)90205-4"},{"key":"e_1_3_2_6_2","doi-asserted-by":"publisher","DOI":"10.1016\/0042-6822(89)90391-7"},{"key":"e_1_3_2_7_2","first-page":"42","volume":"26","year":"2006","unstructured":"Chen, S. L., S. T. Lin, T. C. Tsai, W. C. Hsiao, and Y. P. Tsao. 2006. ErbB4 (JM-b\/CYT-1)-induced expression and phosphorylation of c-Jun is abrogated by human papillomavirus type 16 E5 protein. Oncogene26:42-53.","journal-title":"Oncogene"},{"key":"e_1_3_2_8_2","doi-asserted-by":"publisher","DOI":"10.1128\/jvi.67.10.6170-6178.1993"},{"key":"e_1_3_2_9_2","doi-asserted-by":"publisher","DOI":"10.1006\/excr.1998.4024"},{"key":"e_1_3_2_10_2","doi-asserted-by":"publisher","DOI":"10.1128\/MCB.17.9.5087"},{"key":"e_1_3_2_11_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0042-6822(03)00129-6"},{"key":"e_1_3_2_12_2","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.79.9.5839-5846.2005"},{"key":"e_1_3_2_13_2","doi-asserted-by":"publisher","DOI":"10.1038\/sj.onc.1206554"},{"key":"e_1_3_2_14_2","doi-asserted-by":"publisher","DOI":"10.1158\/0008-5472.CAN-05-0083"},{"key":"e_1_3_2_15_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0042-6822(03)00296-4"},{"key":"e_1_3_2_16_2","doi-asserted-by":"crossref","first-page":"K17","DOI":"10.1016\/j.vaccine.2008.06.021","volume":"26","year":"2008","unstructured":"Giuliano, A. R., G. Tortolero-Luna, E. Ferrer, A. N. Burchell, S. de Sanjose, S. K. Kjaer, N. Mu\u00f1oz, M. Schiffman, and F. X. Bosch. 2008. Epidemiology of human papillomavirus infection in men, cancers other than cervical and benign conditions. Vaccine26S:K17-K28.","journal-title":"Vaccine"},{"key":"e_1_3_2_17_2","doi-asserted-by":"publisher","DOI":"10.1186\/1743-422X-4-116"},{"key":"e_1_3_2_18_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2008.10.011"},{"key":"e_1_3_2_19_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2009.07.034"},{"key":"e_1_3_2_20_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbrc.2008.04.122"},{"key":"e_1_3_2_21_2","doi-asserted-by":"publisher","DOI":"10.2353\/ajpath.2008.080280"},{"key":"e_1_3_2_22_2","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20071311"},{"key":"e_1_3_2_23_2","doi-asserted-by":"publisher","DOI":"10.1186\/1743-422X-5-30"},{"key":"e_1_3_2_24_2","doi-asserted-by":"publisher","DOI":"10.1093\/infdis\/162.6.1263"},{"key":"e_1_3_2_25_2","doi-asserted-by":"publisher","DOI":"10.1006\/meth.1999.0876"},{"key":"e_1_3_2_26_2","doi-asserted-by":"publisher","DOI":"10.1128\/MMBR.68.2.362-372.2004"},{"key":"e_1_3_2_27_2","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.102.2.500"},{"key":"e_1_3_2_28_2","first-page":"57","volume":"7","year":"2007","unstructured":"Miranda, M., and A. Sorkin. 2007. Regulation of receptors and transporters by ubiquitination: new insights into surprisingly similar mechanisms. Mol. Interv.7:57-167.","journal-title":"Mol. Interv."},{"key":"e_1_3_2_29_2","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/18.12.3587"},{"key":"e_1_3_2_30_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.vaccine.2006.05.115"},{"key":"e_1_3_2_31_2","doi-asserted-by":"publisher","DOI":"10.1186\/1475-2867-6-19"},{"key":"e_1_3_2_32_2","first-page":"59","volume":"321","year":"2008","unstructured":"Orth, G. 2008. Host defenses against human papillomaviruses: lessons from epidermodysplasia verruciformis. Curr. Top. Microbiol. Immunol.321:59-83.","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"e_1_3_2_33_2","doi-asserted-by":"publisher","DOI":"10.1016\/0378-1119(92)90589-H"},{"key":"e_1_3_2_34_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.90.18.8392"},{"key":"e_1_3_2_35_2","first-page":"67","year":"2004","unstructured":"Perez, O. D., P. O. Krutzik, and G. P. Nolan. 2004. Flow cytometric analysis of kinase signaling cascade, p. 67-94. In T. S. Hawley and R. G. Hawley (ed.), Flow cytometry protocols, 2nd ed. Humana Press, Totowa, NJ.","journal-title":"Flow cytometry protocols"},{"key":"e_1_3_2_36_2","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.01240-08"},{"key":"e_1_3_2_37_2","doi-asserted-by":"publisher","DOI":"10.1038\/sj.onc.1203718"},{"key":"e_1_3_2_38_2","doi-asserted-by":"publisher","DOI":"10.1128\/jvi.68.9.6120-6123.1994"},{"key":"e_1_3_2_39_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2004.11.011"},{"key":"e_1_3_2_40_2","doi-asserted-by":"publisher","DOI":"10.1038\/sj.onc.1210725"},{"key":"e_1_3_2_41_2","doi-asserted-by":"publisher","DOI":"10.1002\/cyto.990030503"},{"key":"e_1_3_2_42_2","doi-asserted-by":"publisher","DOI":"10.1016\/S1046-5928(03)00049-4"},{"key":"e_1_3_2_43_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.94.9.4451"},{"key":"e_1_3_2_44_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.mi.48.100194.002235"},{"key":"e_1_3_2_45_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2008.11.046"}],"container-title":["Journal of Virology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.asm.org\/doi\/pdf\/10.1128\/JVI.01968-09","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/journals.asm.org\/doi\/pdf\/10.1128\/JVI.01968-09","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,3,5]],"date-time":"2022-03-05T13:48:33Z","timestamp":1646488113000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.asm.org\/doi\/10.1128\/JVI.01968-09"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2010,2,15]]},"references-count":44,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2010,2,15]]}},"alternative-id":["10.1128\/JVI.01968-09"],"URL":"https:\/\/doi.org\/10.1128\/jvi.01968-09","relation":{},"ISSN":["0022-538X","1098-5514"],"issn-type":[{"value":"0022-538X","type":"print"},{"value":"1098-5514","type":"electronic"}],"subject":[],"published":{"date-parts":[[2010,2,15]]},"assertion":[{"value":"2009-09-16","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2009-11-20","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2010-02-15","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}