{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,4]],"date-time":"2026-11-04T02:56:06Z","timestamp":1793760966986,"version":"4.6.1"},"reference-count":90,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,8,20]],"date-time":"2025-08-20T00:00:00Z","timestamp":1755648000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,8,20]],"date-time":"2025-08-20T00:00:00Z","timestamp":1755648000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Social insects employ venom as an external immune defence against pathogens and parasites. Like other Hymenopterans, the venom gland of honey bee (\n                    <jats:italic>Apis mellifera<\/jats:italic>\n                    ) serves as a reservoir of antimicrobial substances, primarily melittin. This study investigated the presence and origin of venom on honey bee body that could act as an external immune defence in honey bee workers infested by the ectoparasite mite,\n                    <jats:italic>Varroa destructor<\/jats:italic>\n                    . Using a multi-step approach, we first confirmed the presence of venom on bees\u2019 bodies using melittin as a marker. We then examined how grooming could facilitate the distribution of venom on the bee\u2019s body through behavioural observations. Further assays were used to compare melittin levels on the bodies of\n                    <jats:italic>Varroa<\/jats:italic>\n                    -free and\n                    <jats:italic>Varroa<\/jats:italic>\n                    -infested workers and assess the effects of bee-venom on mite activity. Our findings confirmed the occurrence of \u201cvenom bathing\u201d in\n                    <jats:italic>A. mellifera<\/jats:italic>\n                    , excluding social components or environmental contamination, with bees likely coating their bodies with antimicrobial substances through self-grooming. Our results further suggest that infested bees spread higher amount of venom on their bodies compared to uninfested bees, and bee-venom significantly reduced mite activity, suggesting that the venom functions as an external defence. However,\n                    <jats:italic>Varroa<\/jats:italic>\n                    negatively impacted melittin production. Our study reveals a previously unknown negative effect of\n                    <jats:italic>V. destructor<\/jats:italic>\n                    : impairment of honey bees\u2019 external immune defence through reduced melittin production.\n                  <\/jats:p>","DOI":"10.1038\/s41598-025-13440-2","type":"journal-article","created":{"date-parts":[[2025,8,20]],"date-time":"2025-08-20T15:45:35Z","timestamp":1755704735000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Varroa destructor weakens the external immunity of western honey bees by impairing melittin production"],"prefix":"10.1038","volume":"15","author":[{"given":"Michelina","family":"Pusceddu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simon","family":"Tragust","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Panagiotis","family":"Theodorou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Irene","family":"Ciabattini Bolla","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jorge","family":"S\u00e1nchez Navarro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francesco","family":"Corrias","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alessandro","family":"Atzei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alberto","family":"Angioni","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ignazio","family":"Floris","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alberto","family":"Satta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,8,20]]},"reference":[{"key":"13440_CR1","doi-asserted-by":"publisher","DOI":"10.1093\/oso\/9780198508175.001.0001","volume-title":"Living in groups","author":"J Krause","year":"2002","unstructured":"Krause, J., Ruxton, G. D. & Ruxton, I. G. Living in groups (Oxford University Press, 2002)."},{"key":"13440_CR2","volume-title":"Parasites in social insects","author":"P Schmid-Hempel","year":"1998","unstructured":"Schmid-Hempel, P. Parasites in social insects Vol. 60 (Princeton University Press, 1998)."},{"issue":"6","key":"13440_CR3","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1016\/j.pt.2017.03.003","volume":"33","author":"P Schmid-Hempel","year":"2017","unstructured":"Schmid-Hempel, P. Parasites and their social hosts. Trends Parasitol. 33(6), 453\u2013462. https:\/\/doi.org\/10.1016\/j.pt.2017.03.003 (2017).","journal-title":"Trends Parasitol."},{"key":"13440_CR4","doi-asserted-by":"publisher","first-page":"R693","DOI":"10.1016\/j.cub.2007.06.008","volume":"17","author":"S Cremer","year":"2007","unstructured":"Cremer, S., Armitage, S. A. O. & Schmid-Hempel, P. Social immunity. Curr. Biol. 17, R693\u2013R702. https:\/\/doi.org\/10.1016\/j.cub.2007.06.008 (2007).","journal-title":"Curr. Biol."},{"key":"13440_CR5","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1146\/annurev-ento-020117-043110","volume":"63","author":"S Cremer","year":"2018","unstructured":"Cremer, S., Pull, C. D. & F\u00fcrst, M. A. Social immunity: emergence and evolution of colony-level disease protection. Annu. Rev. Entomol. 63, 105\u2013123. https:\/\/doi.org\/10.1146\/annurev-ento-020117-043110 (2018).","journal-title":"Annu. Rev. Entomol."},{"key":"13440_CR6","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1146\/annurev.ento.53.103106.093301","volume":"54","author":"N Wilson-Rich","year":"2009","unstructured":"Wilson-Rich, N., Spivak, M., Fefferman, N. H. & Starks, P. T. Genetic, individual, and group facilitation of disease resistance in insect societies. Annu. Rev. Entomol. 54, 405\u2013423. https:\/\/doi.org\/10.1146\/annurev.ento.53.103106.093301 (2009).","journal-title":"Annu. Rev. Entomol."},{"key":"13440_CR7","doi-asserted-by":"publisher","first-page":"S62","DOI":"10.1016\/j.jip.2009.06.019","volume":"103","author":"JD Evans","year":"2010","unstructured":"Evans, J. D. & Spivak, M. Socialized medicine: individual and communal disease barriers in honey bees. J. Invertebr. Pathol. 103, S62\u2013S72. https:\/\/doi.org\/10.1016\/j.jip.2009.06.019 (2010).","journal-title":"J. Invertebr. Pathol."},{"key":"13440_CR8","unstructured":"Shanahan, M. J. Examining propolis use, social immunity, and food systems transformation to support colony health in honey bees and stingless bees. PhD thesis, University of Minnesota (2023)."},{"key":"13440_CR9","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1038\/s41559-023-01969-x","volume":"7","author":"M Stock","year":"2023","unstructured":"Stock, M. et al. Pathogen evasion of social immunity. Nat. Ecol. Evol. 7, 450\u2013460. https:\/\/doi.org\/10.1038\/s41559-023-01969-x (2023).","journal-title":"Nat. Ecol. Evol."},{"key":"13440_CR10","doi-asserted-by":"publisher","first-page":"780","DOI":"10.1016\/j.pt.2024.08.003","volume":"40","author":"S Cremer","year":"2024","unstructured":"Cremer, S. & Pull, C. D. Unconditional versus condition-dependent social immunity. Trends Parasitol. 40, 780\u2013787. https:\/\/doi.org\/10.1016\/j.pt.2024.08.003 (2024).","journal-title":"Trends Parasitol."},{"key":"13440_CR11","doi-asserted-by":"publisher","first-page":"625","DOI":"10.1016\/j.tree.2014.09.002","volume":"29","author":"O Otti","year":"2014","unstructured":"Otti, O., Tragust, S. & Feldhaar, H. Unifying external and internal immune defences. Trends Ecol. Evol. 29, 625\u2013634. https:\/\/doi.org\/10.1016\/j.tree.2014.09.002 (2014).","journal-title":"Trends Ecol. Evol."},{"key":"13440_CR12","doi-asserted-by":"publisher","first-page":"2013","DOI":"10.1098\/rspb.2007.0531","volume":"274","author":"M Chapuisat","year":"2007","unstructured":"Chapuisat, M., Oppliger, A., Magliano, P. & Christe, P. Wood ants use resin to protect themselves against pathogens. Proc. R. Soc. Lond. B Biol. Sci. 274, 2013\u20132017. https:\/\/doi.org\/10.1098\/rspb.2007.0531 (2007).","journal-title":"Proc. R. Soc. Lond. B Biol. Sci."},{"key":"13440_CR13","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1051\/apido\/2010016","volume":"41","author":"M Simone-Finstrom","year":"2010","unstructured":"Simone-Finstrom, M. & Spivak, M. Propolis and bee health: the natural history and significance of resin use by honey bees. Apidologie 41, 295\u2013311. https:\/\/doi.org\/10.1051\/apido\/2010016 (2010).","journal-title":"Apidologie"},{"key":"13440_CR14","doi-asserted-by":"publisher","first-page":"20200003","DOI":"10.1098\/rsbl.2020.0003","volume":"16","author":"P Saelao","year":"2020","unstructured":"Saelao, P., Borba, R. S., Ricigliano, V., Spivak, M. & Simone-Finstrom, M. Honeybee microbiome is stabilized in the presence of propolis. Biol. Lett. 16, 20200003. https:\/\/doi.org\/10.1098\/rsbl.2020.0003 (2020).","journal-title":"Biol. Lett."},{"key":"13440_CR15","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0291744","volume":"19","author":"M Shanahan","year":"2024","unstructured":"Shanahan, M. et al. Thinking inside the box: restoring the propolis envelope facilitates honey bee social immunity. PLoS ONE 19, e0291744. https:\/\/doi.org\/10.1371\/journal.pone.0291744 (2024).","journal-title":"PLoS ONE"},{"key":"13440_CR16","doi-asserted-by":"publisher","first-page":"1591","DOI":"10.1016\/j.anbehav.2007.10.014","volume":"75","author":"G Castella","year":"2008","unstructured":"Castella, G., Chapuisat, M. & Christe, P. Prophylaxis with resin in wood ants. Anim. Behav. 75, 1591\u20131596. https:\/\/doi.org\/10.1016\/j.anbehav.2007.10.014 (2008).","journal-title":"Anim. Behav."},{"key":"13440_CR17","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0034601","volume":"7","author":"MD Simone-Finstrom","year":"2012","unstructured":"Simone-Finstrom, M. D. & Spivak, M. Increased resin collection after parasite challenge: a case of self-medication in honey bees?. PLoS ONE 7, e34601. https:\/\/doi.org\/10.1371\/journal.pone.0034601 (2012).","journal-title":"PLoS ONE"},{"key":"13440_CR18","doi-asserted-by":"publisher","first-page":"3527","DOI":"10.1007\/s00436-018-6050-0","volume":"117","author":"M Pusceddu","year":"2018","unstructured":"Pusceddu, M. et al. The effects of raw propolis on Varroa-infested honey bee (Apis mellifera) workers. Parasitol. Res. 117, 3527\u20133535. https:\/\/doi.org\/10.1007\/s00436-018-6050-0 (2018).","journal-title":"Parasitol. Res."},{"key":"13440_CR19","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1111\/1744-7917.12515","volume":"26","author":"M Pusceddu","year":"2019","unstructured":"Pusceddu, M. et al. Resin foraging dynamics in Varroa destructor infested hives: a case of medication of kin?. Insect Sci. 26, 297\u2013310. https:\/\/doi.org\/10.1111\/1744-7917.12515 (2019).","journal-title":"Insect Sci."},{"key":"13440_CR20","doi-asserted-by":"publisher","first-page":"20212101","DOI":"10.1098\/rspb.2021.2101","volume":"288","author":"M Pusceddu","year":"2021","unstructured":"Pusceddu, M. et al. Honeybees use propolis as a natural pesticide against their major ectoparasite. Proc. R. Soc. Lond. B Biol. Sci. 288, 20212101. https:\/\/doi.org\/10.1098\/rspb.2021.2101 (2021).","journal-title":"Proc. R. Soc. Lond. B Biol. Sci."},{"key":"13440_CR21","doi-asserted-by":"publisher","first-page":"119","DOI":"10.25849\/myrmecol.news_023:119","volume":"23","author":"S Tragust","year":"2016","unstructured":"Tragust, S. External immune defence in ant societies (Hymenoptera: Formicidae): the role of antimicrobial venom and metapleural gland secretion. Myrmecol. News 23, 119\u2013128. https:\/\/doi.org\/10.25849\/myrmecol.news_023:119 (2016).","journal-title":"Myrmecol. News"},{"key":"13440_CR22","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1007\/978-94-007-6458-3_3","volume-title":"Evolution of venomous animals and their toxins","author":"D Baracchi","year":"2017","unstructured":"Baracchi, D. & Tragust, S. Venom as a component of external immune defense in Hymenoptera. In Evolution of venomous animals and their toxins (ed. Malhotra, A.) 213\u2013233 (Dordrecht, Springer, 2017)."},{"key":"13440_CR23","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1146\/annurev.en.27.010182.002011","volume":"27","author":"JO Schmidt","year":"1982","unstructured":"Schmidt, J. O. Biochemistry of insect venoms. Annu. Rev. Entomol. 27, 339\u2013368. https:\/\/doi.org\/10.1146\/annurev.en.27.010182.002011 (1982).","journal-title":"Annu. Rev. Entomol."},{"key":"13440_CR24","doi-asserted-by":"publisher","first-page":"2651","DOI":"10.1007\/s00018-003-3106-8","volume":"60","author":"L Kuhn-Nentwig","year":"2003","unstructured":"Kuhn-Nentwig, L. Antimicrobial and cytolytic peptides of venomous arthropods. Cell. Mol. Life Sci. 60, 2651\u20132668. https:\/\/doi.org\/10.1007\/s00018-003-3106-8 (2003).","journal-title":"Cell. Mol. Life Sci."},{"key":"13440_CR25","doi-asserted-by":"publisher","first-page":"186","DOI":"10.1016\/j.jinsphys.2012.10.005","volume":"59","author":"SJ Moreau","year":"2013","unstructured":"Moreau, S. J. It stings a bit but it cleans well: venoms of Hymenoptera and their antimicrobial potential. J. Insect Physiol. 59, 186\u2013204. https:\/\/doi.org\/10.1016\/j.jinsphys.2012.10.005 (2013).","journal-title":"J. Insect Physiol."},{"key":"13440_CR26","doi-asserted-by":"publisher","first-page":"1","DOI":"10.25849\/myrmecol.news_035:001","volume":"35","author":"L Koch","year":"2025","unstructured":"Koch, L., Niedermeyer, T. H. J. & Tragust, S. Acid reign: Formicine ants and their venoms. Myrmecol. News 35, 1\u201327. https:\/\/doi.org\/10.25849\/myrmecol.news_035:001 (2025).","journal-title":"Myrmecol. News"},{"key":"13440_CR27","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1016\/j.cub.2012.11.034","volume":"23","author":"S Tragust","year":"2013","unstructured":"Tragust, S. et al. Ants disinfect fungus-exposed brood by oral uptake and spread of their poison. Curr. Biol. 23, 76\u201382. https:\/\/doi.org\/10.1016\/j.cub.2012.11.034 (2013).","journal-title":"Curr. Biol."},{"key":"13440_CR28","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1080\/0005772X.1987.11098915","volume":"68","author":"EM Dotimas","year":"1987","unstructured":"Dotimas, E. M. & Hider, R. C. Honeybee venom. Bee World 68, 51\u201370. https:\/\/doi.org\/10.1080\/0005772X.1987.11098915 (1987).","journal-title":"Bee World"},{"key":"13440_CR29","doi-asserted-by":"publisher","first-page":"3360","DOI":"10.3390\/nu12113360","volume":"12","author":"M Carpena","year":"2020","unstructured":"Carpena, M., Nu\u00f1ez-Estevez, B., Soria-Lopez, A. & Simal-Gandara, J. Bee venom: an updating review of its bioactive molecules and its health applications. Nutrients 12, 3360. https:\/\/doi.org\/10.3390\/nu12113360 (2020).","journal-title":"Nutrients"},{"key":"13440_CR30","doi-asserted-by":"publisher","DOI":"10.1016\/j.toxcx.2022.100119","volume":"14","author":"JC Guido-Pati\u00f1o","year":"2022","unstructured":"Guido-Pati\u00f1o, J. C. & Plisson, F. Profiling hymenopteran venom toxins: protein families, structural landscape, biological activities, and pharmacological benefits. Toxicon X 14, 100119. https:\/\/doi.org\/10.1016\/j.toxcx.2022.100119 (2022).","journal-title":"Toxicon X"},{"key":"13440_CR31","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1186\/s12915-023-01656-5","volume":"21","author":"I Koludarov","year":"2023","unstructured":"Koludarov, I. et al. Prevalent bee venom genes evolved before the aculeate stinger and eusociality. BMC Biol. 21, 229. https:\/\/doi.org\/10.1186\/s12915-023-01656-5 (2023).","journal-title":"BMC Biol."},{"key":"13440_CR32","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1016\/j.toxicon.2010.03.023","volume":"56","author":"JRS Ferreira","year":"2010","unstructured":"Ferreira, J. R. S. et al. Africanized honey bee (Apis mellifera) venom profiling: seasonal variation of melittin and phospholipase A2 levels. Toxicon 56, 355\u2013362. https:\/\/doi.org\/10.1016\/j.toxicon.2010.03.023 (2010).","journal-title":"Toxicon"},{"key":"13440_CR33","doi-asserted-by":"publisher","first-page":"1515","DOI":"10.1016\/0022-1910(72)90063-6","volume":"18","author":"H Bachmayer","year":"1972","unstructured":"Bachmayer, H., Kreil, G. & Suchanek, G. Synthesis of promelittin and melittin in the venom gland of queen and worker bees: patterns observed during maturation. J. Insect Physiol. 18, 1515\u20131521. https:\/\/doi.org\/10.1016\/0022-1910(72)90063-6 (1972).","journal-title":"J. Insect Physiol."},{"key":"13440_CR34","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1155\/2012\/258650193","volume":"1","author":"MAASM Ali","year":"2012","unstructured":"Ali, M. A. A. S. M. Studies on bee venom and its medical uses. Int. J. Adv. Res. Technol. 1, 69\u201383. https:\/\/doi.org\/10.1155\/2012\/258650193 (2012).","journal-title":"Int. J. Adv. Res. Technol."},{"key":"13440_CR35","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1016\/B978-0-444-64181-6.00013-9","volume":"60","author":"AA Abd El-Wahed","year":"2019","unstructured":"Abd El-Wahed, A. A. et al. Bee venom composition from chemistry to biological activity. Stud. Nat. Prod. Chem. 60, 459\u2013484. https:\/\/doi.org\/10.1016\/B978-0-444-64181-6.00013-9 (2019).","journal-title":"Stud. Nat. Prod. Chem."},{"key":"13440_CR36","doi-asserted-by":"publisher","first-page":"2090","DOI":"10.3389\/fimmu.2019.02090","volume":"10","author":"MB Pucca","year":"2019","unstructured":"Pucca, M. B. et al. Bee updated: current knowledge on bee venom and bee envenoming therapy. Front. Immunol. 10, 2090. https:\/\/doi.org\/10.3389\/fimmu.2019.02090 (2019).","journal-title":"Front. Immunol."},{"key":"13440_CR37","doi-asserted-by":"publisher","first-page":"917","DOI":"10.1016\/0041-0101(95)00011-a","volume":"33","author":"JO Schmidt","year":"1995","unstructured":"Schmidt, J. O. Toxinology of venoms from the honeybee genus Apis. Toxicon 33, 917\u2013927. https:\/\/doi.org\/10.1016\/0041-0101(95)00011-a (1995).","journal-title":"Toxicon"},{"key":"13440_CR38","doi-asserted-by":"publisher","first-page":"1181","DOI":"10.1016\/0041-0101(95)00054-p","volume":"33","author":"MD Owen","year":"1995","unstructured":"Owen, M. D. & Pfaff, L. A. Melittin synthesis in the venom system of the honey bee (Apis mellifera L.). Toxicon 33, 1181\u20131188. https:\/\/doi.org\/10.1016\/0041-0101(95)00054-p (1995).","journal-title":"Toxicon"},{"key":"13440_CR39","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1590\/S1678-91992003000200002","volume":"9","author":"PR De Lima","year":"2003","unstructured":"De Lima, P. R. & Brochetto-Braga, M. R. Hymenoptera venom review focusing on Apis mellifera. J. Venom Anim. Toxins Incl. Trop. Dis. 9, 149\u2013162. https:\/\/doi.org\/10.1590\/S1678-91992003000200002 (2003).","journal-title":"J. Venom Anim. Toxins Incl. Trop. Dis."},{"key":"13440_CR40","doi-asserted-by":"publisher","first-page":"2867","DOI":"10.2174\/1381612828666220928110103","volume":"28","author":"T Kontogiannis","year":"2022","unstructured":"Kontogiannis, T., Dimitriou, T. G., Didaras, N. A. & Mossialos, D. Antiviral activity of bee products. Curr. Pharm. Des. 28, 2867\u20132878. https:\/\/doi.org\/10.2174\/1381612828666220928110103 (2022).","journal-title":"Curr. Pharm. Des."},{"key":"13440_CR41","doi-asserted-by":"publisher","first-page":"314","DOI":"10.1126\/science.177.4046.314","volume":"177","author":"E Habermann","year":"1972","unstructured":"Habermann, E. Bee and wasp venoms: the biochemistry and pharmacology of their peptides and enzymes are reviewed. Science 177, 314\u2013322. https:\/\/doi.org\/10.1126\/science.177.4046.314 (1972).","journal-title":"Science"},{"key":"13440_CR42","doi-asserted-by":"publisher","first-page":"1793","DOI":"10.1096\/fasebj.12.15.1793","volume":"12","author":"O Froy","year":"1998","unstructured":"Froy, O. & Gurevitz, M. Membrane potential modulators: a thread of scarlet from plants to humans. FASEB J. 12, 1793\u20131796. https:\/\/doi.org\/10.1096\/fasebj.12.15.1793 (1998).","journal-title":"FASEB J."},{"key":"13440_CR43","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1016\/j.jinsphys.2009.11.013","volume":"56","author":"D Baracchi","year":"2010","unstructured":"Baracchi, D. & Turillazzi, S. Differences in venom and cuticular peptides in individuals of Apis mellifera (Hymenoptera: Apidae) determined by MALDI-TOF MS. J. Insect Physiol. 56, 366\u2013375. https:\/\/doi.org\/10.1016\/j.jinsphys.2009.11.013 (2010).","journal-title":"J. Insect Physiol."},{"key":"13440_CR44","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1016\/j.toxicon.2011.08.017","volume":"58","author":"D Baracchi","year":"2011","unstructured":"Baracchi, D., Francese, S. & Turillazzi, S. Beyond the antipredatory defence: honey bee venom function as a component of social immunity. Toxicon 58, 550\u2013557. https:\/\/doi.org\/10.1016\/j.toxicon.2011.08.017 (2011).","journal-title":"Toxicon"},{"key":"13440_CR45","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1080\/00218839.2016.1196016","volume":"55","author":"DJ Pritchard","year":"2016","unstructured":"Pritchard, D. J. Grooming by honey bees as a component of Varroa resistant behavior. J. Apic. Res. 55, 38\u201348. https:\/\/doi.org\/10.1080\/00218839.2016.1196016 (2016).","journal-title":"J. Apic. Res."},{"key":"13440_CR46","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1007\/BF01218573","volume":"16","author":"R B\u00fcchler","year":"1992","unstructured":"B\u00fcchler, R., Drescher, W. & Tornier, I. Grooming behaviour of Apis cerana, Apis mellifera and Apis dorsata and its effect on the parasitic mites Varroa jacobsoni and Tropilaelaps clareae. Exp. Appl. Acarol. 16, 313\u2013319. https:\/\/doi.org\/10.1007\/BF01218573 (1992).","journal-title":"Exp. Appl. Acarol."},{"key":"13440_CR47","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1051\/apido:19960101","volume":"27","author":"I Fries","year":"1996","unstructured":"Fries, I., Huazhen, W., Wei, S. & Jin, C. S. Grooming behavior and damaged mites (Varroa jacobsoni) in Apis cerana cerana and Apis mellifera ligustica. Apidologie 27, 3\u201311. https:\/\/doi.org\/10.1051\/apido:19960101 (1996).","journal-title":"Apidologie"},{"key":"13440_CR48","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1051\/APIDO:19980304","volume":"29","author":"JS Pettis","year":"1998","unstructured":"Pettis, J. S. & Pankiw, T. Grooming behavior by Apis mellifera L. in the presence of Acarapis woodi (Rennie) (Acari: Tarsonemidae). Apidologie 29, 241\u2013253. https:\/\/doi.org\/10.1051\/APIDO:19980304 (1998).","journal-title":"Apidologie"},{"key":"13440_CR49","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1051\/apido:19990205","volume":"30","author":"O Boecking","year":"1999","unstructured":"Boecking, O. & Spivak, M. Behavioral defenses of honey bees against Varroa jacobsoni Oud. Apidologie 30, 141\u2013158. https:\/\/doi.org\/10.1051\/apido:19990205 (1999).","journal-title":"Apidologie"},{"key":"13440_CR50","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1080\/00218839.2016.1159055","volume":"54","author":"C Invernizzi","year":"2022","unstructured":"Invernizzi, C., Zefferino, I., Santos, E., S\u00e1nchez, L. & Mendoza, Y. Multilevel assessment of grooming behavior against Varroa destructor in Italian and Africanized honey bees. J. Apic. Res. 54, 321\u2013327. https:\/\/doi.org\/10.1080\/00218839.2016.1159055 (2022).","journal-title":"J. Apic. Res."},{"key":"13440_CR51","doi-asserted-by":"publisher","first-page":"766","DOI":"10.1186\/1471-2164-14-766","volume":"14","author":"R Li","year":"2013","unstructured":"Li, R. et al. Proteome and phosphoproteome analysis of honeybee (Apis mellifera) venom collected from electrical stimulation and manual extraction of the venom gland. BMC Genomics 14, 766. https:\/\/doi.org\/10.1186\/1471-2164-14-766 (2013).","journal-title":"BMC Genomics"},{"key":"13440_CR52","unstructured":"SANTE\/11312\/2021. Analytical quality control and method validation procedures for pesticide residues analysis in food and feed SANTE 11312\/2021"},{"key":"13440_CR53","doi-asserted-by":"publisher","first-page":"3265","DOI":"10.1007\/s00216-023-04705-8","volume":"415","author":"A Beasley-Green","year":"2023","unstructured":"Beasley-Green, A. & Heckert, N. A. Estimation of measurement uncertainty for the quantification of protein by ID-LC\u2013MS\/MS. Anal. Bioanal. Chem. 415, 3265\u20133274. https:\/\/doi.org\/10.1007\/s00216-023-04705-8 (2023).","journal-title":"Anal. Bioanal. Chem."},{"key":"13440_CR54","unstructured":"Commission of the European Communities. In Pappas, N. & Thrasyvoulou, A. (eds) Searching for an accurate method to evaluate the degree of Varroa infestation in honeybee colonies, 1st edn, 85\u201392 (European Research on Varroatosis Control, Rotterdam, 1988)."},{"key":"13440_CR55","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3896\/IBRA.1.52.1.04","volume":"52","author":"GR Williams","year":"2013","unstructured":"Williams, G. R. et al. Standard methods for maintaining adult Apis mellifera in cages under in vitro laboratory conditions. J. Apic. Res. 52, 1\u201336. https:\/\/doi.org\/10.3896\/IBRA.1.52.1.04 (2013).","journal-title":"J. Apic. Res."},{"key":"13440_CR56","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1002735","volume":"8","author":"F Nazzi","year":"2012","unstructured":"Nazzi, F. et al. Synergistic parasite-pathogen interactions mediated by host immunity can drive the collapse of honeybee colonies. PLoS Pathog. 8, e1002735. https:\/\/doi.org\/10.1371\/journal.ppat.1002735 (2012).","journal-title":"PLoS Pathog."},{"key":"13440_CR57","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1163\/156853974X00534","volume":"49","author":"J Altmann","year":"1974","unstructured":"Altmann, J. Observational study of behaviour: sampling methods. Behaviour 49, 227\u2013265. https:\/\/doi.org\/10.1163\/156853974X00534 (1974).","journal-title":"Behaviour"},{"key":"13440_CR58","doi-asserted-by":"publisher","first-page":"415","DOI":"10.1051\/apido:19950507","volume":"26","author":"N Milani","year":"1995","unstructured":"Milani, N. The resistance of Varroa jacobsoni Oud to pyrethroids: a laboratory assay. Apidologie 26, 415\u2013429. https:\/\/doi.org\/10.1051\/apido:19950507 (1995).","journal-title":"Apidologie"},{"key":"13440_CR59","unstructured":"R Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria (2018). https:\/\/www.R-project.org\/"},{"key":"13440_CR60","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v067.i01","volume":"67","author":"D Bates","year":"2015","unstructured":"Bates, D., Maechler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1\u201348. https:\/\/doi.org\/10.18637\/jss.v067.i01 (2015).","journal-title":"J. Stat. Softw."},{"key":"13440_CR61","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1002\/bimj.200810425","volume":"50","author":"T Hothorn","year":"2008","unstructured":"Hothorn, T., Bretz, F. & Westfall, P. Simultaneous inference in general parametric models. Biometr. J. 50, 346\u2013363. https:\/\/doi.org\/10.1002\/bimj.200810425 (2008).","journal-title":"Biometr. J."},{"key":"13440_CR62","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1111\/j.1439-0310.2011.01895.x","volume":"117","author":"TL Stout","year":"2011","unstructured":"Stout, T. L., Slone, J. D. & Schneider, S. S. Age and behavior of honey bee workers, Apis mellifera, that interact with drones. Ethology 117, 459\u2013468. https:\/\/doi.org\/10.1111\/j.1439-0310.2011.01895.x (2011).","journal-title":"Ethology"},{"key":"13440_CR63","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1007\/s00040-013-0310-x","volume":"60","author":"A Goins","year":"2013","unstructured":"Goins, A. & Schneider, S. S. Drone, quality and caste interactions in the honey bee. Apis mellifera L. Insectes Soc. 60, 453\u2013461. https:\/\/doi.org\/10.1007\/s00040-013-0310-x (2013).","journal-title":"Apis mellifera L. Insectes Soc."},{"key":"13440_CR64","doi-asserted-by":"publisher","first-page":"1687","DOI":"10.1007\/s00265-015-1980-3","volume":"69","author":"C Tranter","year":"2015","unstructured":"Tranter, C. & Hughes, W. H. O. Acid, silk and grooming: alternative strategies in social immunity in ants?. Behav. Ecol. Sociobiol. 69, 1687\u20131699. https:\/\/doi.org\/10.1007\/s00265-015-1980-3 (2015).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"13440_CR65","doi-asserted-by":"publisher","first-page":"97","DOI":"10.21203\/rs.3.rs-3964303\/v1","volume":"78","author":"NI Beydizada","year":"2024","unstructured":"Beydizada, N. I., Abels, A., Schultheiss, P. & Frank, E. T. Injury-dependent wound care behavior in the desert ant Cataglyphis nodus. Behav. Ecol. Sociobiol. 78, 97. https:\/\/doi.org\/10.21203\/rs.3.rs-3964303\/v1 (2024).","journal-title":"Behav. Ecol. Sociobiol."},{"key":"13440_CR66","doi-asserted-by":"publisher","unstructured":"Baumann, K. Evolution of the venom system in aculeate hymenoptera. Thesis, The University of Queensland, School of Biological Sciences (2018). https:\/\/doi.org\/10.14264\/uql.2018.752","DOI":"10.14264\/uql.2018.752"},{"key":"13440_CR67","doi-asserted-by":"publisher","first-page":"S96","DOI":"10.1016\/j.jip.2009.07.016","volume":"103","author":"P Rosenkranz","year":"2010","unstructured":"Rosenkranz, P., Aumeier, P. & Ziegelmann, B. Biology and control of Varroa destructor. J. Invertebr. Pathol. 103, S96\u2013S119. https:\/\/doi.org\/10.1016\/j.jip.2009.07.016 (2010).","journal-title":"J. Invertebr. Pathol."},{"key":"13440_CR68","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1146\/annurev-ento-010715-023731","volume":"61","author":"F Nazzi","year":"2016","unstructured":"Nazzi, F. & Le Conte, Y. Ecology of Varroa destructor, the major ectoparasite of the western honey bee. Apis mellifera. Annu. Rev. Entomol. 61, 417\u2013432. https:\/\/doi.org\/10.1146\/annurev-ento-010715-023731 (2016).","journal-title":"Apis mellifera. Annu. Rev. Entomol."},{"key":"13440_CR69","doi-asserted-by":"publisher","first-page":"eabj1398","DOI":"10.1126\/sciadv.abj1398","volume":"7","author":"M Pusceddu","year":"2021","unstructured":"Pusceddu, M. et al. Honey bees increase social distancing when facing the ectoparasite Varroa destructor. Sci. Adv. 7, eabj1398. https:\/\/doi.org\/10.1126\/sciadv.abj1398 (2021).","journal-title":"Sci. Adv."},{"key":"13440_CR70","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/0022-2011(87)90125-X","volume":"49","author":"YS Peng","year":"1987","unstructured":"Peng, Y. S., Fang, Y., Xu, S. & Ge, L. The resistance mechanism of the Asian honey bee, Apis cerana Fabr., to an ectoparasitic mite. Varroa jacobsoni Oudemans. J. Invertebr. Pathol. 49, 54\u201360. https:\/\/doi.org\/10.1016\/0022-2011(87)90125-X (1987).","journal-title":"Varroa jacobsoni Oudemans. J. Invertebr. Pathol."},{"key":"13440_CR71","doi-asserted-by":"publisher","first-page":"8928","DOI":"10.1038\/s41598-020-65780-w","volume":"10","author":"A Cini","year":"2020","unstructured":"Cini, A. et al. Increased immunocompetence and network centrality of allogroomer workers suggest a link between individual and social immunity in honeybees. Sci. Rep. 10, 8928. https:\/\/doi.org\/10.1038\/s41598-020-65780-w (2020).","journal-title":"Sci. Rep."},{"key":"13440_CR72","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1051\/apido:2001006","volume":"33","author":"A Garedew","year":"2002","unstructured":"Garedew, A., Lamprecht, I., Schmolz, E. & Schricker, B. The varroacidal action of propolis: a laboratory assay. Apidologie 33, 41\u201350. https:\/\/doi.org\/10.1051\/apido:2001006 (2002).","journal-title":"Apidologie"},{"key":"13440_CR73","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1007\/s13592-014-0289-y","volume":"45","author":"NJ Fern\u00e1ndez","year":"2014","unstructured":"Fern\u00e1ndez, N. J. et al. A scientific note on the first report of honeybee venom inhibiting Paenibacillus larvae growth. Apidologie 45, 719\u2013721. https:\/\/doi.org\/10.1007\/s13592-014-0289-y (2014).","journal-title":"Apidologie"},{"key":"13440_CR74","doi-asserted-by":"publisher","DOI":"10.1590\/1678-9199-JVATITD-2022-0025","author":"AA Sani","year":"2022","unstructured":"Sani, A. A. et al. Inhibitory activities of propolis, nisin, melittin and essential oil compounds on Paenibacillus alvei and Bacillus subtilis. J. Venom. Anim. Toxins Incl. Trop. Dis https:\/\/doi.org\/10.1590\/1678-9199-JVATITD-2022-0025 (2022).","journal-title":"J. Venom. Anim. Toxins Incl. Trop. Dis"},{"key":"13440_CR75","doi-asserted-by":"publisher","first-page":"943","DOI":"10.3389\/fmicb.2020.00943","volume":"11","author":"O Ya\u00f1ez","year":"2020","unstructured":"Ya\u00f1ez, O. et al. Bee viruses: routes of infection in Hymenoptera. Front. Microbiol. 11, 943. https:\/\/doi.org\/10.3389\/fmicb.2020.00943 (2020).","journal-title":"Front. Microbiol."},{"key":"13440_CR76","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1080\/00218839.1983.11100586","volume":"22","author":"L Bailey","year":"1983","unstructured":"Bailey, L., Ball, B. V. & Perry, J. N. Honeybee paralysis: its natural spread and its diminished incidence in England and Wales. J. Apic. Res. 22, 191\u2013195. https:\/\/doi.org\/10.1080\/00218839.1983.11100586 (1983).","journal-title":"J. Apic. Res."},{"key":"13440_CR77","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1016\/j.pestbp.2017.10.009","volume":"144","author":"M Coulon","year":"2018","unstructured":"Coulon, M. et al. Metabolisation of thiamethoxam (a neonicotinoid pesticide) and interaction with the Chronic bee paralysis virus in honeybees. Pestic. Biochem. Physiol. 144, 10\u201318. https:\/\/doi.org\/10.1016\/j.pestbp.2017.10.009 (2018).","journal-title":"Pestic. Biochem. Physiol."},{"key":"13440_CR78","doi-asserted-by":"publisher","first-page":"1188","DOI":"10.3390\/v6031188","volume":"6","author":"E Amiri","year":"2014","unstructured":"Amiri, E., Meixner, M., B\u00fcchler, R. & Kryger, P. Chronic bee paralysis virus in honeybee queens: evaluating susceptibility and infection routes. Viruses 6, 1188\u20131201. https:\/\/doi.org\/10.3390\/v6031188 (2014).","journal-title":"Viruses"},{"key":"13440_CR79","doi-asserted-by":"publisher","first-page":"9","DOI":"10.3390\/insects10010009","volume":"10","author":"E Amiri","year":"2019","unstructured":"Amiri, E. et al. Israeli acute paralysis virus: honey bee queen\u2013worker interaction and potential virus transmission pathways. Insects 10, 9. https:\/\/doi.org\/10.3390\/insects10010009 (2019).","journal-title":"Insects"},{"key":"13440_CR80","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s41938-022-00580-0","volume":"32","author":"H Seyam","year":"2022","unstructured":"Seyam, H. et al. Effect of honeybee venom and Egyptian propolis on the honeybee (Apis mellifera L.) health in vivo. Egypt J Biol Pest Control 32, 1\u201310. https:\/\/doi.org\/10.1186\/s41938-022-00580-0 (2022).","journal-title":"Egypt J Biol Pest Control"},{"key":"13440_CR81","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1007\/s13592-023-01048-2","volume":"55","author":"SH Mahmoud","year":"2024","unstructured":"Mahmoud, S. H., Kandel, M., El-Seedi, H. & Al Naggar, Y. Honey bee venom promotes the immune system and reduces Vairimorpha (Nosema) ceranae infection in honey bees (Apis mellifera L.). Apidologie 55, 8. https:\/\/doi.org\/10.1007\/s13592-023-01048-2 (2024).","journal-title":"Apidologie"},{"key":"13440_CR82","doi-asserted-by":"publisher","DOI":"10.1016\/j.cbpc.2024.109928","volume":"281","author":"M Hejn\u00edkov\u00e1","year":"2024","unstructured":"Hejn\u00edkov\u00e1, M., Tom\u010dala, A., \u010cern\u00fd, J. & Kodr\u00edk, D. Melittin\u2014The principal toxin of honeybee venom\u2014Is also produced in the honeybee fat body. Comp Biochem Physiol C Toxicol Pharmacol 281, 109928. https:\/\/doi.org\/10.1016\/j.cbpc.2024.109928 (2024).","journal-title":"Comp Biochem Physiol C Toxicol Pharmacol"},{"key":"13440_CR83","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1007\/s13592-022-00977-8","volume":"54","author":"I Grindrod","year":"2023","unstructured":"Grindrod, I. & Martin, S. J. Varroa resistance in Apis cerana: a review. Apidologie 54, 14. https:\/\/doi.org\/10.1007\/s13592-022-00977-8 (2023).","journal-title":"Apidologie"},{"key":"13440_CR84","doi-asserted-by":"publisher","first-page":"506","DOI":"10.3390\/antiox10040506","volume":"10","author":"I Floris","year":"2021","unstructured":"Floris, I., Pusceddu, M. & Satta, A. The Sardinian bitter honey: from ancient healing use to recent findings. Antioxidants 10, 506. https:\/\/doi.org\/10.3390\/antiox10040506 (2021).","journal-title":"Antioxidants"},{"key":"13440_CR85","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1016\/j.fbio.2019.04.004","volume":"29","author":"A Felicioli","year":"2019","unstructured":"Felicioli, A. et al. In vitro antibacterial activity and volatile characterisation of organic Apis mellifera ligustica (Spinola, 1906) beeswax ethanol extracts. Food Biosci 29, 102\u2013109. https:\/\/doi.org\/10.1016\/j.fbio.2019.04.004 (2019).","journal-title":"Food Biosci"},{"key":"13440_CR86","doi-asserted-by":"publisher","first-page":"811","DOI":"10.3390\/antibiotics9110811","volume":"9","author":"NA Didaras","year":"2020","unstructured":"Didaras, N. A., Karatasou, K., Dimitriou, T. G., Amoutzias, G. D. & Mossialos, D. Antimicrobial activity of bee-collected pollen and beebread: state of the art and future perspectives. Antibiotics 9, 811. https:\/\/doi.org\/10.3390\/antibiotics9110811 (2020).","journal-title":"Antibiotics"},{"key":"13440_CR87","doi-asserted-by":"publisher","first-page":"50","DOI":"10.2174\/2211352513666150318234430","volume":"13","author":"MF Alreshoodi","year":"2015","unstructured":"Alreshoodi, M. F. & Sultanbawa, Y. Antimicrobial activity of royal jelly. Anti-Infect. Agents 13, 50\u201359. https:\/\/doi.org\/10.2174\/2211352513666150318234430 (2015).","journal-title":"Anti-Infect. Agents"},{"key":"13440_CR88","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1007\/s13592-015-0400-z","volume":"47","author":"S Erler","year":"2016","unstructured":"Erler, S. & Moritz, R. F. A. Pharmacophagy and pharmacophory: mechanisms of self-medication and disease prevention in the honeybee colony (Apis mellifera). Apidologie 47, 389\u2013411. https:\/\/doi.org\/10.1007\/s13592-015-0400-z (2016).","journal-title":"Apidologie"},{"key":"13440_CR89","doi-asserted-by":"publisher","first-page":"338","DOI":"10.1016\/j.pt.2024.02.003","volume":"40","author":"S Erler","year":"2024","unstructured":"Erler, S. et al. Insects\u2019 essential role in understanding and broadening animal medication. Trends Parasitol 40, 338\u2013349. https:\/\/doi.org\/10.1016\/j.pt.2024.02.003 (2024).","journal-title":"Trends Parasitol"},{"key":"13440_CR90","doi-asserted-by":"publisher","first-page":"2249","DOI":"10.1002\/ece3.2834","volume":"7","author":"T Br\u00fctsch","year":"2017","unstructured":"Br\u00fctsch, T., Jaffuel, G., Vallat, A., Turlings, T. C. & Chapuisat, M. Wood ants produce a potent antimicrobial agent by applying formic acid on tree-collected resin. Ecol Evol 7, 2249\u20132254. https:\/\/doi.org\/10.1002\/ece3.2834 (2017).","journal-title":"Ecol Evol"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-13440-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-13440-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-13440-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,9]],"date-time":"2025-09-09T09:08:12Z","timestamp":1757408892000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-13440-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,20]]},"references-count":90,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["13440"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-13440-2","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-6238801\/v1","asserted-by":"object"}]},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,20]]},"assertion":[{"value":"16 March 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 July 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"20 August 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"No ethical approval is required for this study.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}}],"article-number":"30623"}}