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While investigating a potential link between RHBDL4-regulated p24 (TMED) family proteins and proTGF\u03b1 trafficking, we uncovered an RHBDL4-independent pathway in which the cargo receptor TMED9 induces unconventional protein secretion (UcPS) of ectopically expressed proTGF\u03b1. This pathway selectively requires GRASP65 (also known as GORASP1), but not the closely related GRASP55 (or GORASP2), and depends on components of the autophagic machinery and the ESCRT-associated protein ALIX (also known as PDCD6IP), consistent with a Golgi-bypass secretion route. TMED9- and GRASP65-induced proTGF\u03b1 secretion is insensitive to brefeldin A treatment and involves trafficking through pre-Golgi compartments. Although largely based on overexpression systems, our findings identify a previously unrecognized UcPS mechanism and provide insight into alternative protein trafficking pathways that might be exploited under conditions of proteostasis stress and in pathological contexts such as cancer and inflammation. At present, our data establish the existence of a TMED9- and GRASP65-dependent Golgi-bypass trafficking route under conditions of ectopic cargo expression, whereas its endogenous physiological relevance remains to be established.<\/jats:p>","DOI":"10.1242\/jcs.264719","type":"journal-article","created":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T08:29:01Z","timestamp":1785140941000},"source":"Crossref","is-referenced-by-count":0,"title":["A Golgi-bypass secretion mechanism for proTGF\u03b1 involving TMED9 and GRASP65"],"prefix":"10.1242","volume":"139","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-9322-5916","authenticated-orcid":false,"given":"Susanne S.","family":"Steigleder","sequence":"first","affiliation":[{"name":"Center for Biochemistry and Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD) 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]},{"name":"University of Cologne 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Isabel","family":"D\u00f6ring","sequence":"additional","affiliation":[{"name":"Center for Biochemistry and Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD) 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]},{"name":"University of Cologne 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marina","family":"Tauber","sequence":"additional","affiliation":[{"name":"Center for Biochemistry and Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD) 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]},{"name":"University of Cologne 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8126-415X","authenticated-orcid":false,"given":"Julian","family":"N\u00fcchel","sequence":"additional","affiliation":[{"name":"Center for Biochemistry and Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD) 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]},{"name":"University of Cologne 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]},{"name":"Center for Molecular Medicine Cologne (CMMC) 2 , Faculty of Medicine and , 50931 Cologne ,","place":["Germany"]},{"name":"University Hospital Cologne, University of Cologne 2 , Faculty of Medicine and , 50931 Cologne ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0996-1268","authenticated-orcid":false,"given":"Marius K.","family":"Lemberg","sequence":"additional","affiliation":[{"name":"Center for Biochemistry and Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD) 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]},{"name":"University of Cologne 1 , Faculty of Medicine , , 50931 Cologne ,","place":["Germany"]},{"name":"Center for Molecular Medicine Cologne (CMMC) 2 , Faculty of Medicine and , 50931 Cologne ,","place":["Germany"]},{"name":"University Hospital Cologne, University of Cologne 2 , Faculty of Medicine and , 50931 Cologne ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,8,28]]},"reference":[{"key":"2026082806574947700_JCS264719C1","doi-asserted-by":"publisher","first-page":"102219","DOI":"10.1016\/j.jbc.2022.102219","article-title":"GRASP55 regulates the unconventional secretion and aggregation of mutant huntingtin","volume":"298","author":"Ahat","year":"2022","journal-title":"J. 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