{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,10]],"date-time":"2026-01-10T00:10:39Z","timestamp":1768003839312,"version":"3.49.0"},"reference-count":87,"publisher":"SAGE Publications","issue":"4","license":[{"start":{"date-parts":[[2023,4,1]],"date-time":"2023-04-01T00:00:00Z","timestamp":1680307200000},"content-version":"unspecified","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"name":"UMIB\/FCT","award":["UIDB\/00215\/2020"],"award-info":[{"award-number":["UIDB\/00215\/2020"]}]},{"name":"UMIB\/FCT","award":["UIDP\/00215\/2020"],"award-info":[{"award-number":["UIDP\/00215\/2020"]}]},{"name":"ITR\/FCT","award":["LA\/P\/0064\/2020"],"award-info":[{"award-number":["LA\/P\/0064\/2020"]}]}],"content-domain":{"domain":["journals.sagepub.com"],"crossmark-restriction":true},"short-container-title":["J Histochem Cytochem."],"published-print":{"date-parts":[[2023,4]]},"abstract":"<jats:p> Coronavirus disease (COVID-19) is caused by severe acute respiratory syndrome coronavirus-2 (SARS-CoV-2). SARS-CoV-2 RNA has been found in the human testis on occasion, but subgenomic SARS-CoV-2 and infectious SARS-CoV-2 virions have not been found. There is no direct evidence of SARS-CoV-2 infection of testicular cells. To better understand this, it is necessary to determine whether SARS-CoV-2 receptors and proteases are present in testicular cells. To overcome this limitation, we focused on elucidating with immunohistochemistry the spatial distribution of the SARS-CoV-2 receptors angiotensin-converting enzyme 2 (ACE2) and cluster of differentiation 147 (CD147), as well as their viral spike protein priming proteases, transmembrane protease serine 2 (TMPRSS2) and cathepsin L (CTSL), required for viral fusion with host cells. At the protein level, human testicular tissue expressed both receptors and proteases studied. Both ACE2 and TMPRSS2 were found in interstitial cells (endothelium, Leydig, and myoid peritubular cells) and in the seminiferous epithelium (Sertoli cells, spermatogonia, spermatocytes, and spermatids). The presence of CD147 was observed in all cell types except endothelium and peritubular cells, while CTSL was exclusively observed in Leydig, peritubular, and Sertoli cells. These findings show that the ACE2 receptor and its protease TMPRSS2 are coexpressed in all testicular cells, as well as the CD147 receptor and its protease CTSL in Leydig and Sertoli cells, indicating that testicular SARS-CoV-2 infection cannot be ruled out without further investigation: <\/jats:p>","DOI":"10.1369\/00221554231168916","type":"journal-article","created":{"date-parts":[[2023,4,7]],"date-time":"2023-04-07T08:57:22Z","timestamp":1680857842000},"page":"169-197","update-policy":"https:\/\/doi.org\/10.1177\/sage-journals-update-policy","source":"Crossref","is-referenced-by-count":9,"title":["Spatial Distribution of SARS-CoV-2 Receptors and Proteases in Testicular Cells"],"prefix":"10.1369","volume":"71","author":[{"given":"Maria Rita","family":"Ribeiro","sequence":"first","affiliation":[{"name":"Laboratory of Cell Biology, Department of Microscopy, ICBAS \u2013 School of Medicine and Biomedical Sciences, UMIB \u2013 Unit for Multidisciplinary Research in Biomedicine \/ ITR \u2013 Laboratory for Integrative and Translational Research in Population Health, University of Porto, Porto, Portugal"},{"name":"Department of Veterinary Sciences, School of Agricultural and Veterinary Sciences (ECAV), CECAV \u2013 Interdisciplinary Research Center in Animal Health, Associate Laboratory for Animal and Veterinary Science (AL4AnimalS), University of Tr\u00e1s-os-Montes and Alto Douro (UTAD), Vila Real, Portugal"}]},{"given":"Ana Margarida","family":"Calado","sequence":"additional","affiliation":[{"name":"Department of Veterinary Sciences, School of Agricultural and Veterinary Sciences (ECAV), CECAV \u2013 Interdisciplinary Research Center in Animal Health, Associate Laboratory for Animal and Veterinary Science (AL4AnimalS), University of Tr\u00e1s-os-Montes and Alto Douro (UTAD), Vila Real, Portugal"}]},{"given":"\u00c2ngela","family":"Alves","sequence":"additional","affiliation":[{"name":"Laboratory of Cell Biology, Department of Microscopy, ICBAS \u2013 School of Medicine and Biomedical Sciences, UMIB \u2013 Unit for Multidisciplinary Research in Biomedicine \/ ITR \u2013 Laboratory for Integrative and Translational Research in Population Health, University of Porto, Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8434-5086","authenticated-orcid":false,"given":"Rute","family":"Pereira","sequence":"additional","affiliation":[{"name":"Laboratory of Cell Biology, Department of Microscopy, ICBAS \u2013 School of Medicine and Biomedical Sciences, UMIB \u2013 Unit for Multidisciplinary Research in Biomedicine \/ ITR \u2013 Laboratory for Integrative and Translational Research in Population Health, University of Porto, Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3009-3290","authenticated-orcid":false,"given":"M\u00e1rio","family":"Sousa","sequence":"additional","affiliation":[{"name":"Laboratory of Cell Biology, Department of Microscopy, ICBAS \u2013 School of Medicine and Biomedical Sciences, UMIB \u2013 Unit for Multidisciplinary Research in Biomedicine \/ ITR \u2013 Laboratory for Integrative and Translational Research in Population Health, University of Porto, Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6551-3822","authenticated-orcid":false,"given":"Ros\u00e1lia","family":"S\u00e1","sequence":"additional","affiliation":[{"name":"Laboratory of Cell Biology, Department of Microscopy, ICBAS \u2013 School of Medicine and Biomedical Sciences, UMIB \u2013 Unit for Multidisciplinary Research in Biomedicine \/ ITR \u2013 Laboratory for Integrative and Translational Research in Population Health, University of Porto, Porto, Portugal"},{"name":"Faculty of Medicine, University of Porto, Port, Portugal"}]}],"member":"179","published-online":{"date-parts":[[2023,4,7]]},"reference":[{"key":"bibr1-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(20)30566-3"},{"key":"bibr2-00221554231168916","unstructured":"World Health Organization (WHO). Novel coronavirus (2019-nCoV): situation report, 1. Geneva, Switzerland: World Health Organization; 2020."},{"key":"bibr3-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2020.10.004"},{"key":"bibr4-00221554231168916","unstructured":"Centers for Disease Control and Prevention (CDC). https:\/\/www.cdc.gov\/coronavirus\/2019-ncov\/faq.html. Accessed June 1, 2022."},{"key":"bibr5-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/s40618-020-01261-1"},{"key":"bibr6-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/s40618-021-01580-x"},{"key":"bibr7-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1111\/andr.12939"},{"key":"bibr8-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1111\/andr.12848"},{"key":"bibr9-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1093\/biolre\/ioaa050"},{"key":"bibr10-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.fertnstert.2020.05.028"},{"key":"bibr11-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1159\/000510531"},{"key":"bibr12-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1038\/s41443-020-00344-0"},{"key":"bibr13-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1111\/and.13912"},{"key":"bibr14-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1001\/jamanetworkopen.2020.8292"},{"key":"bibr15-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1056\/NEJMoa2001191"},{"issue":"18","key":"bibr16-00221554231168916","volume":"323","author":"Wang W","year":"2020","journal-title":"JAMA"},{"key":"bibr17-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1093\/humrep\/deab026"},{"key":"bibr18-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/s10815-021-02119-y"},{"issue":"4","key":"bibr19-00221554231168916","volume":"5","author":"Gorbalenya AE","year":"2020","journal-title":"Nat Microbiol"},{"key":"bibr20-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbadis.2020.165878"},{"key":"bibr21-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4939-2438-7_1"},{"key":"bibr22-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2020.01818"},{"key":"bibr23-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1038\/s41423-021-00752-2"},{"key":"bibr24-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-020-2180-5"},{"key":"bibr25-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.03.045"},{"key":"bibr26-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.tibs.2021.06.001"},{"key":"bibr27-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3390\/v12050497"},{"key":"bibr28-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-021-00418-x"},{"key":"bibr29-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1101\/2020.03.14.988345."},{"key":"bibr30-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2020.589505"},{"key":"bibr31-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.02.052"},{"key":"bibr32-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.antiviral.2020.104792"},{"key":"bibr33-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.heliyon.2022.e08799"},{"key":"bibr34-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmi.2020.04.023"},{"key":"bibr35-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/s11684-020-0754-0"},{"key":"bibr36-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2020.108175"},{"key":"bibr37-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3390\/cells9040920"},{"key":"bibr38-00221554231168916","doi-asserted-by":"publisher","DOI":"10.15252\/msb.20209610"},{"key":"bibr39-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1093\/oxfordjournals.humrep.a019004"},{"key":"bibr40-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1093\/humrep\/12.1.80"},{"key":"bibr41-00221554231168916","volume":"16","author":"Cooper TG","year":"2010","journal-title":"Hum Reprod Update"},{"key":"bibr42-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1152\/japplphysiol.00808.2006"},{"key":"bibr43-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-80464-1"},{"key":"bibr44-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1161\/CIRCRESAHA.120.317015"},{"key":"bibr45-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2020.559841"},{"key":"bibr46-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3390\/microorganisms8101492"},{"key":"bibr47-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21217943"},{"key":"bibr48-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.fertnstert.2015.11.004"},{"key":"bibr49-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.mehy.2020.110287"},{"key":"bibr50-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1093\/biolre\/ioaa080"},{"key":"bibr51-00221554231168916","doi-asserted-by":"publisher","DOI":"10.2147\/IDR.S270543"},{"key":"bibr52-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.fertnstert.2020.05.001"},{"key":"bibr53-00221554231168916","doi-asserted-by":"publisher","DOI":"10.20944\/preprints202002.0307.v1."},{"key":"bibr54-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1101\/2020.04.06.028522."},{"key":"bibr55-00221554231168916","doi-asserted-by":"publisher","DOI":"10.3389\/fmed.2020.563893"},{"key":"bibr56-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1136\/jclinpath-2020-206987"},{"key":"bibr57-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/s11427-020-1705-0"},{"key":"bibr58-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1038\/s41392-020-00426-x"},{"key":"bibr59-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.semcdb.2014.02.008"},{"key":"bibr60-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.omtn.2021.11.009"},{"key":"bibr61-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1093\/molehr\/gar013"},{"key":"bibr62-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/s11033-018-4568-y"},{"key":"bibr63-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.lfs.2020.118313"},{"key":"bibr64-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijantimicag.2020.106004"},{"key":"bibr65-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1006\/scdb.1998.0203"},{"key":"bibr66-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ph.50.030188.002411"},{"key":"bibr67-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1679\/aohc.59.1"},{"issue":"1","key":"bibr68-00221554231168916","volume":"93","author":"Fave RFD","year":"2021","journal-title":"Arch Ital Urol Androl"},{"key":"bibr69-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.11.032"},{"key":"bibr70-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-020-2179-y"},{"key":"bibr71-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.virusres.2014.11.021"},{"key":"bibr72-00221554231168916","doi-asserted-by":"publisher","DOI":"10.7150\/ijms.46695"},{"key":"bibr73-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1080\/13685538.2020.1807930"},{"key":"bibr74-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1007\/s12020-020-02515-x"},{"key":"bibr75-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1111\/andr.12821"},{"key":"bibr76-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1101\/2020.03.21.20037267."},{"key":"bibr77-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1080\/22221751.2021.1969869"},{"key":"bibr78-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1210\/jcem-7-7-493"},{"key":"bibr79-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/S0022-5347(17)58673-5"},{"key":"bibr80-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1111\/jcmm.15541"},{"key":"bibr81-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.fertnstert.2020.12.033"},{"key":"bibr82-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1128\/MMBR.65.2.208-231.2001"},{"issue":"4","key":"bibr83-00221554231168916","first-page":"84","volume":"16","author":"Chen WK","year":"2020","journal-title":"J Mens Health"},{"key":"bibr84-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(20)30937-5"},{"key":"bibr85-00221554231168916","doi-asserted-by":"publisher","DOI":"10.2174\/0929867322666150821094334"},{"key":"bibr86-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1183\/13993003.01028-2020"},{"key":"bibr87-00221554231168916","doi-asserted-by":"publisher","DOI":"10.1016\/j.tox.2010.07.003"}],"container-title":["Journal of Histochemistry &amp; Cytochemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.sagepub.com\/doi\/pdf\/10.1369\/00221554231168916","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/journals.sagepub.com\/doi\/full-xml\/10.1369\/00221554231168916","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/journals.sagepub.com\/doi\/pdf\/10.1369\/00221554231168916","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,2,28]],"date-time":"2025-02-28T19:19:05Z","timestamp":1740770345000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.sagepub.com\/doi\/10.1369\/00221554231168916"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,4]]},"references-count":87,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2023,4]]}},"alternative-id":["10.1369\/00221554231168916"],"URL":"https:\/\/doi.org\/10.1369\/00221554231168916","relation":{},"ISSN":["0022-1554","1551-5044"],"issn-type":[{"value":"0022-1554","type":"print"},{"value":"1551-5044","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,4]]}}}