{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,18]],"date-time":"2026-07-18T09:40:31Z","timestamp":1784367631295,"version":"3.55.0"},"reference-count":206,"publisher":"Portland Press Ltd.","issue":"03","license":[{"start":{"date-parts":[[2025,5,14]],"date-time":"2025-05-14T00:00:00Z","timestamp":1747180800000},"content-version":"vor","delay-in-days":20,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["portlandpress.com"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2025,6,30]]},"abstract":"<jats:p>Intercellular communication is an essential hallmark of multicellular organisms for their development and adult tissue homeostasis. Over the past two decades, attention has been focused on communication mechanisms based on various membrane structures, as illustrated by the burst of scientific literature in the field of extracellular vesicles (EVs). These lipid bilayer-bound nano- or microparticles, as vehicle-like devices, act as regulators in various biological and physiological processes. When EVs are internalized by recipient cells, their membrane and cytoplasmic cargoes can interfere with cellular activities, affecting pathways that regulate cell proliferation, differentiation, and migration. In cancer, EVs can transfer oncogenic factors, stimulate neo-angiogenesis and immunosuppression, reprogram stromal cells, and confer drug resistance traits, thereby remodeling the surrounding microenvironment. Although the mechanisms underlying EV biogenesis and uptake are now better understood, little is known about the spatiotemporal mechanism(s) of their actions after internalization. In this respect, we have shown that a fraction of endocytosed EVs reaches the nuclear compartment via the VOR (VAP-A-ORP3-Rab7) complex-mediated docking of late endosomes to the outer nuclear membrane in the nucleoplasmic reticulum, positioning and facilitating the transfer of EV cargoes into the nucleoplasm via nuclear pores. Here, we highlight the EV heterogeneity, the cellular pathways governing EV release and uptake by donor and recipient cells, respectively, and focus on a novel intracellular pathway leading to the nuclear transfer of EV cargoes. We will discuss how to intercept it, which could open up new avenues for clinical applications in which EVs and other small extracellular particles (e.g., retroviruses) are implicated.<\/jats:p>","DOI":"10.1042\/bst20253005","type":"journal-article","created":{"date-parts":[[2025,5,14]],"date-time":"2025-05-14T14:22:41Z","timestamp":1747232561000},"page":"529-546","update-policy":"https:\/\/doi.org\/10.1042\/crossmark_policy","source":"Crossref","is-referenced-by-count":8,"title":["Extracellular membrane particles <i>en route<\/i> to the nucleus \u2013 exploring the VOR complex"],"prefix":"10.1042","volume":"53","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0644-7375","authenticated-orcid":false,"given":"Aurelio","family":"Lorico","sequence":"first","affiliation":[{"name":"1Department of Basic Sciences, College of Osteopathic Medicine, Touro University Nevada, Henderson, NV 89014, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6347-8576","authenticated-orcid":false,"given":"Mark F.","family":"Santos","sequence":"additional","affiliation":[{"name":"1Department of Basic Sciences, College of Osteopathic Medicine, Touro University Nevada, Henderson, NV 89014, U.S.A."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7687-0983","authenticated-orcid":false,"given":"Jana","family":"Karbanov\u00e1","sequence":"additional","affiliation":[{"name":"2Biotechnology Center (BIOTEC), Center for Molecular and Cellular Bioengineering, Technische Universit\u00e4t Dresden, Dresden, Saxony, Germany"},{"name":"3Tissue Engineering Laboratories, Medizinische Fakult\u00e4t der Technischen Universit\u00e4t Dresden, Dresden, Saxony, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1181-3659","authenticated-orcid":false,"given":"Denis","family":"Corbeil","sequence":"additional","affiliation":[{"name":"2Biotechnology Center (BIOTEC), Center for Molecular and Cellular Bioengineering, Technische Universit\u00e4t Dresden, Dresden, Saxony, Germany"},{"name":"3Tissue Engineering Laboratories, Medizinische Fakult\u00e4t der Technischen Universit\u00e4t Dresden, Dresden, Saxony, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"288","published-online":{"date-parts":[[2025,4,24]]},"reference":[{"key":"2025063011412567100_BST-2025-3005C1","doi-asserted-by":"crossref","DOI":"10.1002\/jev2.12404","article-title":"Minimal information for studies of extracellular vesicles (MISEV2023): From basic to advanced approaches","volume":"13","author":"Welsh","year":"2024","journal-title":"J. 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