{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T17:41:25Z","timestamp":1773337285653,"version":"3.50.1"},"reference-count":60,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T00:00:00Z","timestamp":1773273600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell. Infect. Microbiol."],"abstract":"<jats:p>\n                    Since the onset of COVID-19, emphasis has been largely placed on spike proteins, while other molecules of severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), such as the 78-aa-long envelope protein (E-pr), have been largely overlooked. This study was conducted to identify the orientation of E-pr in the envelope of the living SARS-CoV-2 virus and to confirm whether the viroid E-pr plays a role in the attachment and stimulation of the host cells. Using a customized antisera against the C-terminal half of E-pr (EC\n                    <jats:sub>38<\/jats:sub>\n                    , 38 aa), we demonstrated that the EC\n                    <jats:sub>38<\/jats:sub>\n                    segment was located outside of the intact virus. Synthetic peptides with the sequence of the EC\n                    <jats:sub>38<\/jats:sub>\n                    segment were bound to and modulated gene expression in cultured human vascular endothelial cells (HUVECs), and altered genes were enriched for the ontology terms \u201chemostasis\u201d and \u201cregulation of systemic arterial blood pressure by hormone.\u201d EC\n                    <jats:sub>38<\/jats:sub>\n                    -mediated pull-down of HUVEC membrane proteins plus mass spectrometry revealed that the EC\n                    <jats:sub>38<\/jats:sub>\n                    -interacting proteins were enriched for \u201cactin cytoskeleton organization,\u201d \u201ccell\u2013cell interaction,\u201d \u201ccell\u2013cell adhesion,\u201d \u201chemostasis,\u201d \u201cviral infection pathways,\u201d \u201ccytokine signaling in immune system,\u201d etc., among which CKAP4, vimentin, and ATP5B were further identified to bind EC\n                    <jats:sub>38<\/jats:sub>\n                    peptides. Molecular docking supported the potential binding of the E-pr with the ATP synthase. In summary, by displaying a C-terminal-outward orientation on the surface of the SARS-CoV-2 virus, E-pr was able to adhere to and stimulate host cells. Future studies should address whether the E-pr pathway could be utilized as a target for managing SARS-CoV-2 infection in humans.\n                  <\/jats:p>","DOI":"10.3389\/fcimb.2026.1776252","type":"journal-article","created":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T06:50:14Z","timestamp":1773298214000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["C-terminus-outward orientation of SARS-CoV-2 envelope proteins on viral capsid enables a novel virus\u2013cell interaction pathway"],"prefix":"10.3389","volume":"16","author":[{"given":"Jie","family":"Xu","sequence":"first","affiliation":[{"name":"Eye Institute of Xiamen University, School of Medicine, Xiamen University","place":["Xiamen, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wei","family":"Zhao","sequence":"additional","affiliation":[{"name":"Eye Institute of Xiamen University, School of Medicine, Xiamen University","place":["Xiamen, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuanyuan","family":"Li","sequence":"additional","affiliation":[{"name":"Department of Laboratory Examination, People\u2019s Hospital of Rizhao City","place":["Rizhao, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jing","family":"Zheng","sequence":"additional","affiliation":[{"name":"Xiamen Center for Disease Control and Prevention","place":["Xiamen, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yulian","family":"Wang","sequence":"additional","affiliation":[{"name":"Eye Institute of Xiamen University, School of Medicine, Xiamen University","place":["Xiamen, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Huimin","family":"Sun","sequence":"additional","affiliation":[{"name":"Xiang\u2019an Hospital of Xiamen University, Xiamen University","place":["Xiamen, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sijin","family":"Wu","sequence":"additional","affiliation":[{"name":"Wisdom Lake Academy of Pharmacy, Xi\u2019an Jiaotong-Liverpool University","place":["Suzhou, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Baoqing","family":"Fu","sequence":"additional","affiliation":[{"name":"Xiang\u2019an Hospital of Xiamen University, Xiamen University","place":["Xiamen, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yiqiang","family":"Wang","sequence":"additional","affiliation":[{"name":"Eye Institute of Xiamen University, School of Medicine, Xiamen University","place":["Xiamen, China"]},{"name":"Xiang\u2019an Hospital of Xiamen University, Xiamen University","place":["Xiamen, China"]},{"name":"Wisdom Lake Academy of Pharmacy, Xi\u2019an Jiaotong-Liverpool University","place":["Suzhou, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2026,3,12]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2020.00405","article-title":"Functional pangenome analysis shows key features of E protein are preserved in SARS and SARS-CoV-2","volume":"10","author":"Alam","year":"2020","journal-title":"Front. 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