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The main limitation is their dependence on freshly drawn blood, which limits batch production. Long-term storage of EDTA-anticoagulated blood at \u221280\u00b0C without cryoprotectant represents an accessible alternative, but its effect on EMV physicochemical properties and protein composition has not been investigated. We prepared EMVs from fresh and frozen blood and characterized the resulting fractions by dynamic light scattering, nano-flow cytometry, cryo-transmission electron microscopy, negative staining transmission electron microscopy, and quantitative mass spectrometry-based proteomics. EMV size, surface charge, and morphology did not differ significantly between fresh and frozen preparations. Extrusion effectively normalized particle size despite increased heterogeneity in frozen ghost cell fractions, and both preparations yielded structurally intact vesicles after ultracentrifugation. At the proteomic level, freezing reduced proteomic complexity in membrane fractions and selectively depleted cytosol-associated proteins. The core membrane proteome, including Band 3, remained stable. However, glycophorin A showed the largest reduction in frozen preparations, and CD47 levels were reduced in the ultracentrifuged fraction. These differences may be critical for applications where preservation of specific surface proteins is required, such as immune evasion or targeted drug delivery. In some cases, frozen blood is potentially a viable source for EMV production when immediate processing is not feasible.<\/jats:p>","DOI":"10.1021\/acs.jproteome.6c00535","type":"journal-article","created":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T07:01:57Z","timestamp":1788937317000},"page":"5534-5548","source":"Crossref","is-referenced-by-count":0,"title":["Proteomic and\nPhysicochemical Analysis of Erythrocyte\nMembrane Vesicles from Fresh and Frozen Blood"],"prefix":"10.1021","volume":"25","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-2393-5674","authenticated-orcid":true,"given":"Katarina","family":"Tomic\u0301","sequence":"first","affiliation":[{"name":"Joz\u030cef Stefan Institute , , ,","place":["Ljubljana, Slovenia, 1000"]},{"name":"Joz\u030cef Stefan International Postgraduate School, Joz\u030cef Stefan Institute , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sara","family":"Ivanovski","sequence":"additional","affiliation":[{"name":"Joz\u030cef Stefan International Postgraduate School, Joz\u030cef Stefan Institute , ,","place":["Ljubljana, Slovenia, 1000"]},{"name":"Joz\u030cef Stefan Institute , , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jernej","family":"S\u030cribar","sequence":"additional","affiliation":[{"name":"Joz\u030cef Stefan Institute , , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-6702-1266","authenticated-orcid":true,"given":"Das\u030ca","family":"C\u030cebulj","sequence":"additional","affiliation":[{"name":"Joz\u030cef Stefan Institute , , ,","place":["Ljubljana, Slovenia, 1000"]},{"name":"University of Ljubljana , , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samuel","family":"Z\u030cvanut","sequence":"additional","affiliation":[{"name":"University of Ljubljana , , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Metka","family":"Lenassi","sequence":"additional","affiliation":[{"name":"University of Ljubljana , , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8375-6713","authenticated-orcid":true,"given":"Marko","family":"Fonovic\u0301","sequence":"additional","affiliation":[{"name":"Joz\u030cef Stefan Institute , , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0203-0708","authenticated-orcid":true,"given":"Igor","family":"Kriz\u030caj","sequence":"additional","affiliation":[{"name":"Joz\u030cef Stefan Institute , , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6551-0250","authenticated-orcid":true,"given":"Nina","family":"Kostevs\u030cek","sequence":"additional","affiliation":[{"name":"Joz\u030cef Stefan Institute , , ,","place":["Ljubljana, Slovenia, 1000"]},{"name":"Joz\u030cef Stefan International Postgraduate School, Joz\u030cef Stefan Institute , ,","place":["Ljubljana, Slovenia, 1000"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"issue":"2","key":"2026100208421626100_cit1","doi-asserted-by":"publisher","first-page":"695","DOI":"10.3390\/pharmaceutics15020695","article-title":"Drug Delivery Systems in Regenerative Medicine: An\nUpdated Review","volume":"15","author":"Mansour","year":"2023","journal-title":"Pharmaceutics"},{"issue":"13","key":"2026100208421626100_cit2","doi-asserted-by":"publisher","first-page":"1989","DOI":"10.3390\/cells11131989","article-title":"Different Sourced Extracellular Vesicles\nand Their Potential Applications\nin Clinical Treatments","volume":"11","author":"Bahmani","year":"2022","journal-title":"Cells"},{"key":"2026100208421626100_cit3","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1016\/j.addr.2015.09.012","article-title":"PEGylation as a\nStrategy for Improving Nanoparticle-Based Drug and Gene Delivery","volume":"99","author":"Suk","year":"2016","journal-title":"Adv. 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Biopharm."},{"issue":"9","key":"2026100208421626100_cit30","doi-asserted-by":"publisher","first-page":"941","DOI":"10.1038\/nbt.3330","article-title":"Principles\nof Nanoparticle Design\nfor Overcoming Biological Barriers to Drug Delivery","volume":"33","author":"Blanco","year":"2015","journal-title":"Nat. Biotechnol."},{"issue":"32","key":"2026100208421626100_cit31","doi-asserted-by":"publisher","first-page":"2301706","DOI":"10.1002\/adhm.202301706","article-title":"Assessing\nExtracellular Vesicles in Human Biofluids Using Flow-Based Analyzers","volume":"12","author":"Yim","year":"2023","journal-title":"Adv. Healthc. Mater."},{"issue":"11","key":"2026100208421626100_cit32","doi-asserted-by":"publisher","DOI":"10.1002\/jex2.107","article-title":"Human Red Blood Cells Release Microvesicles with Distinct\nSizes and Protein Composition That Alter Neutrophil Phagocytosis","volume":"2","author":"De Oliveira","year":"2023","journal-title":"J. Extracell. 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Extracell. Vesicles"},{"issue":"6","key":"2026100208421626100_cit39","doi-asserted-by":"publisher","first-page":"603","DOI":"10.3390\/biomedicines9060603","article-title":"Perspectives\nof Microscopy Methods for Morphology Characterisation of Extracellular\nVesicles from Human Biofluids","volume":"9","author":"Malenica","year":"2021","journal-title":"Biomedicines"},{"issue":"1","key":"2026100208421626100_cit40","doi-asserted-by":"publisher","first-page":"1535750","DOI":"10.1080\/20013078.2018.1535750","article-title":"Minimal Information for Studies of Extracellular Vesicles\n2018 (MISEV2018): A Position Statement of the International Society\nfor Extracellular Vesicles and Update of the MISEV2014 Guidelines","volume":"7","author":"The\u0301ry","year":"2018","journal-title":"J. Extracell. Vesicles"},{"issue":"3","key":"2026100208421626100_cit41","doi-asserted-by":"publisher","first-page":"396","DOI":"10.1016\/j.jprot.2009.07.010","article-title":"Comparative Proteomics\nof Erythrocyte Aging in Vivo and in Vitro","volume":"73","author":"Bosman","year":"2010","journal-title":"J.\nProteomics"},{"issue":"1","key":"2026100208421626100_cit42","doi-asserted-by":"publisher","first-page":"206","DOI":"10.3390\/life13010206","article-title":"Impact of Experimental Conditions\non Extracellular Vesicles\u2019 Proteome: A Comparative Study","volume":"13","author":"Bo\u0308ro\u0308czky","year":"2023","journal-title":"Life"},{"key":"2026100208421626100_cit43","doi-asserted-by":"publisher","first-page":"587936","DOI":"10.1101\/2024.04.03.587936","article-title":"Extracellular Vesicles Isolated from\nFrozen and Fresh Human Melanoma Tissue Are Similar in Purity and Protein\nComposition","author":"D\u2019Arrigo","year":"2024","journal-title":"bioarxiv"},{"issue":"24","key":"2026100208421626100_cit44","doi-asserted-by":"publisher","first-page":"12969","DOI":"10.1073\/pnas.94.24.12969","article-title":"Erythrocyte Membrane\nVesiculation: Model for the Molecular Mechanism of Protein Sorting","volume":"94","author":"Knowles","year":"1997","journal-title":"Proc. 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