{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T19:28:19Z","timestamp":1791487699562,"version":"4.3.4"},"reference-count":52,"publisher":"MDPI AG","issue":"10","license":[{"start":{"date-parts":[[2025,10,6]],"date-time":"2025-10-06T00:00:00Z","timestamp":1759708800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100007155","name":"Medical Research Council","doi-asserted-by":"publisher","award":["MR\/W019353\/1"],"award-info":[{"award-number":["MR\/W019353\/1"]}],"id":[{"id":"10.13039\/501100007155","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Toxins"],"abstract":"<jats:p>The puff adder (Bitis arietans) is a clinically relevant viper species found throughout Africa, and it is responsible for a greater incidence of health-related envenomations than all other snake species on the continent combined. Unresolved skeletal muscle damage is a common consequence of B. arietans envenomation that can result in long-term morbidity and even death. Antivenom treatment can mitigate the systemic effects of the venom but offers little protection against local tissue damage. Identifying the mechanisms through which B. arietans venom induces tissue damage and impedes skeletal muscle regeneration could identify possible treatment alternatives that could help alleviate the long-term consequences of envenomation. Skeletal muscle has an innate ability to regenerate, but constituents within the venom can impede multiple stages of this regeneration process. In this study, we employed a combination of biochemical analyses, cell-based assays, and in vivo experiments to assess the toxicological implications of B. arietans envenomation and its impacts on key processes of regeneration. Our findings demonstrate that the pathological characteristics of permanent muscle damage resulting from B. arietans envenomation may be attributed to the venom\u2019s effects on muscle stem cell precursors, the extracellular matrix (ECM), and the influence of blood-borne proteins that promote fibrosis.<\/jats:p>","DOI":"10.3390\/toxins17100496","type":"journal-article","created":{"date-parts":[[2025,10,6]],"date-time":"2025-10-06T13:33:41Z","timestamp":1759757621000},"page":"496","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Compromised Regeneration, Damage to Blood Vessels and the Endomysium Underpin Permanent Muscle Damage Induced by Puff Adder (Bitis arietans) Venom"],"prefix":"10.3390","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7664-7340","authenticated-orcid":false,"given":"Sodiq Opeyemi","family":"Adeyemi","sequence":"first","affiliation":[{"name":"School of Biological Sciences, University of Reading, Reading RG6 6UB, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-5900-5226","authenticated-orcid":false,"given":"Nicholas John","family":"Richards","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Reading, Reading RG6 6UB, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ali","family":"Alqallaf","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Reading, Reading RG6 6UB, UK"},{"name":"Medical Services Authority, Ministry of Defence, Kuwait City 13012, Kuwait"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Husain Bin","family":"Haidar","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Reading, Reading RG6 6UB, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0227-6564","authenticated-orcid":false,"given":"Mustafa Jawad Jalil","family":"Al-Asady","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Reading, Reading RG6 6UB, UK"},{"name":"Ministry of Higher Education and Scientific Research, Baghdad 8CH9+3JH, Iraq"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4647-1399","authenticated-orcid":false,"given":"Jarred","family":"Williams","sequence":"additional","affiliation":[{"name":"School of Pharmacy, University of Reading, Reading RG6 6UB, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4637-4468","authenticated-orcid":false,"given":"Jos\u00e9 R.","family":"Almeida","sequence":"additional","affiliation":[{"name":"School of Pharmacy, University of Reading, Reading RG6 6UB, UK"},{"name":"Biomolecules Discovery Group, Universidad Regional Amaz\u00f3nica Ikiam, Tena 150150, Ecuador"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ketan","family":"Patel","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Reading, Reading RG6 6UB, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,10,6]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1186\/s41182-025-00754-0","article-title":"Toward achieving the WHO 2030 target in the mitigation of snakebite envenoming: The fundamental challenges in addressing the unmet needs","volume":"53","author":"Allen","year":"2025","journal-title":"Trop. Med. Health"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"100074","DOI":"10.1016\/j.toxcx.2021.100074","article-title":"Chronic health effects and cost of snakebite","volume":"9\u201310","author":"Kasturiratne","year":"2021","journal-title":"Toxicon X"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1186\/s40409-017-0127-6","article-title":"Snakebite envenomation turns again into a neglected tropical disease!","volume":"23","author":"Chippaux","year":"2017","journal-title":"J. Venom. Anim. Toxins Incl. Trop. Dis."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"100076","DOI":"10.1016\/j.toxcx.2021.100076","article-title":"Addressing the global snakebite crisis with geo-spatial analyses\u2014Recent advances and future direction","volume":"11","author":"Pintor","year":"2021","journal-title":"Toxicon X"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"100086","DOI":"10.1016\/j.toxcx.2021.100086","article-title":"Access to antivenoms in the developing world: A multidisciplinary analysis","volume":"12","author":"Potet","year":"2021","journal-title":"Toxicon X"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/S0140-6736(09)61754-2","article-title":"Snake bite","volume":"375","author":"Warrell","year":"2010","journal-title":"Lancet"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1186\/s41182-024-00581-9","article-title":"Toxicological analyses of the venoms of Nigerian vipers Echis ocellatus and Bitis arietans","volume":"52","author":"Dingwoke","year":"2024","journal-title":"Trop. Med. Health"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1186\/1678-9199-19-27","article-title":"Public health aspects of snakebite care in West Africa: Perspectives from Nigeria","volume":"19","author":"Habib","year":"2013","journal-title":"J. Venom. Anim. Toxins Incl. Trop. Dis."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"911","DOI":"10.1081\/CLT-120016963","article-title":"Severe puff adder (Bitis arietans) envenomation with coagulopathy","volume":"40","author":"Lavonas","year":"2002","journal-title":"J. Toxicol. Clin. Toxicol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1186\/s40409-018-0176-5","article-title":"Kn-Ba: A novel serine protease isolated from Bitis arietans snake venom with fibrinogenolytic and kinin-releasing activities","volume":"24","author":"Megale","year":"2018","journal-title":"J. Venom. Anim. Toxins Incl. Trop. Dis."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Youngman, N.J., Debono, J., Dobson, J.S., Zdenek, C.N., Harris, R.J., Op den Brouw, B., Coimbra, F.C.P., Naude, A., Coster, K., and Sundman, E. (2019). Venomous Landmines: Clinical Implications of Extreme Coagulotoxic Diversification and Differential Neutralization by Antivenom of Venoms within the Viperid Snake Genus Bitis. Toxins, 11.","DOI":"10.3390\/toxins11070422"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1016\/0041-0101(86)90037-1","article-title":"Amount of venom injected by Russell's viper (Vipera russelli)","volume":"24","author":"Tun","year":"1986","journal-title":"Toxicon"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e02330","DOI":"10.1002\/eap.2330","article-title":"Proximity between humans and a highly medically significant snake, Russell's viper, in a tropical rural community","volume":"31","author":"Glaudas","year":"2021","journal-title":"Ecol. Appl."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"915","DOI":"10.1016\/j.toxicon.2003.11.005","article-title":"Skeletal muscle degeneration induced by venom phospholipases A2: Insights into the mechanisms of local and systemic myotoxicity","volume":"42","author":"Ownby","year":"2003","journal-title":"Toxicon"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Hern\u00e1ndez-Altamirano, J.A., Salazar-Valenzuela, D., Medina-Villamizar, E.J., Quirola, D.R., Patel, K., Vaiyapuri, S., Lomonte, B., and Almeida, J.R. (2022). First Insights into the Venom Composition of Two Ecuadorian Coral Snakes. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms232314686"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1016\/0041-0101(85)90367-8","article-title":"Myonecrosis, myoglobinuria and acute renal failure induced by South American rattlesnake (Crotalus durissus terrificus) envenomation in Brazil","volume":"23","author":"Cupo","year":"1985","journal-title":"Toxicon"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Guti\u00e9rrez, J.M., Escalante, T., Hern\u00e1ndez, R., Gastaldello, S., Saravia-Otten, P., and Rucavado, A. (2018). Why is Skeletal Muscle Regeneration Impaired after Myonecrosis Induced by Viperid Snake Venoms?. Toxins, 10.","DOI":"10.3390\/toxins10050182"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"6094","DOI":"10.1038\/s41467-020-19981-6","article-title":"A therapeutic combination of two small molecule toxin inhibitors provides broad preclinical efficacy against viper snakebite","volume":"11","author":"Albulescu","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Hern\u00e1ndez, R., Cabalceta, C., Saravia-Otten, P., Chaves, A., Guti\u00e9rrez, J.M., and Rucavado, A. (2011). Poor Regenerative Outcome after Skeletal Muscle Necrosis Induced by Bothrops asper Venom: Alterations in Microvasculature and Nerves. PLoS ONE, 6.","DOI":"10.1371\/annotation\/9caa2be8-b5e6-4553-8575-f0b575442172"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Forcina, L., Cosentino, M., and Musar\u00f2, A. (2020). Mechanisms Regulating Muscle Regeneration: Insights into the Interrelated and Time-Dependent Phases of Tissue Healing. Cells, 9.","DOI":"10.3390\/cells9051297"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Musar\u00f2, A. (2014). The Basis of Muscle Regeneration. Adv. Biol., 2014.","DOI":"10.1155\/2014\/612471"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1111\/j.1469-7580.2009.01138.x","article-title":"The origin, molecular regulation and therapeutic potential of myogenic stem cell populations","volume":"215","author":"Otto","year":"2009","journal-title":"J. Anat."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Williams, H.F., Mellows, B.A., Mitchell, R., Sfyri, P., Layfield, H.J., Salamah, M., Vaiyapuri, R., Collins-Hooper, H., Bicknell, A.B., and Matsakas, A. (2019). Mechanisms underpinning the permanent muscle damage induced by snake venom metalloprotease. PLoS Negl. Trop. Dis., 13.","DOI":"10.1371\/journal.pntd.0007041"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Bin Haidar, H., Almeida, J.R., Williams, J., Guo, B., Bigot, A., Senthilkumaran, S., Vaiyapuri, S., and Patel, K. (2024). Differential effects of the venoms of Russell's viper and Indian cobra on human myoblasts. Sci. Rep., 14.","DOI":"10.1038\/s41598-024-53366-9"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1007\/s10974-023-09662-4","article-title":"Alterations of the skeletal muscle contractile apparatus in necrosis induced by myotoxic snake venom phospholipases A(2): A mini-review","volume":"45","author":"Lomonte","year":"2024","journal-title":"J. Muscle Res. Cell Motil."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Sonavane, M., Almeida, J.R., Rajan, E., Williams, H.F., Townsend, F., Cornish, E., Mitchell, R.D., Patel, K., and Vaiyapuri, S. (2023). Intramuscular Bleeding and Formation of Microthrombi during Skeletal Muscle Damage Caused by a Snake Venom Metalloprotease and a Cardiotoxin. Toxins, 15.","DOI":"10.3390\/toxins15090530"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1016\/S0300-9084(00)01163-9","article-title":"Snake venom metalloproteinases: Their role in the pathogenesis of local tissue damage","volume":"82","author":"Rucavado","year":"2000","journal-title":"Biochimie"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1597","DOI":"10.1111\/febs.16693","article-title":"Regenerative inflammation: When immune cells help to re-build tissues","volume":"291","author":"Nagy","year":"2024","journal-title":"Febs. J."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1457","DOI":"10.1016\/j.devcel.2024.03.017","article-title":"Three-dimensional imaging studies in mice identify cellular dynamics of skeletal muscle regeneration","volume":"59","author":"Collins","year":"2024","journal-title":"Dev Cell"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Howard, E.E., Pasiakos, S.M., Blesso, C.N., Fussell, M.A., and Rodriguez, N.R. (2020). Divergent Roles of Inflammation in Skeletal Muscle Recovery From Injury. Front. Physiol., 11.","DOI":"10.3389\/fphys.2020.00087"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.semcdb.2021.05.005","article-title":"Myeloid cell diversification during regenerative inflammation: Lessons from skeletal muscle","volume":"119","author":"Patsalos","year":"2021","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Rodr\u00edguez, C., Tim\u00f3teo-Ferreira, F., Minchiotti, G., Brunelli, S., and Guardiola, O. (2024). Cellular interactions and microenvironment dynamics in skeletal muscle regeneration and disease. Front. Cell Dev. Biol., 12.","DOI":"10.3389\/fcell.2024.1385399"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Wang, X., and Zhou, L. (2022). The Many Roles of Macrophages in Skeletal Muscle Injury and Repair. Front. Cell Dev. Biol., 10.","DOI":"10.3389\/fcell.2022.952249"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"7058","DOI":"10.1038\/s41467-022-34191-y","article-title":"Depletion of CD206(+) M2-like macrophages induces fibro-adipogenic progenitors activation and muscle regeneration","volume":"13","author":"Nawaz","year":"2022","journal-title":"Nat. Commun."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1641","DOI":"10.1101\/gad.247205.114","article-title":"Myomaker is essential for muscle regeneration","volume":"28","author":"Millay","year":"2014","journal-title":"Genes Dev."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.yexcr.2004.08.004","article-title":"Desmin: A major intermediate filament protein essential for the structural integrity and function of muscle","volume":"301","author":"Paulin","year":"2004","journal-title":"Exp Cell Res"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Waiddyanatha, S., Silva, A., Weerakoon, K., Siribaddana, S., and Isbister, G.K. (2022). Long-term health effects perceived by snakebite patients in rural Sri Lanka: A cohort study. PLOS Neglected Trop. Dis., 16.","DOI":"10.1371\/journal.pntd.0010723"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Harrison, R.A., and Guti\u00e9rrez, J.M. (2016). Priority Actions and Progress to Substantially and Sustainably Reduce the Mortality, Morbidity and Socioeconomic Burden of Tropical Snakebite. Toxins, 8.","DOI":"10.3390\/toxins8120351"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1007\/s00204-022-03420-0","article-title":"A current perspective on snake venom composition and constituent protein families","volume":"97","author":"Tasoulis","year":"2023","journal-title":"Arch. Toxicol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1016\/0014-4800(91)90002-F","article-title":"Skeletal muscle degeneration and regeneration after injection of bothropstoxin-II, a phospholipase A2 isolated from the venom of the snake Bothrops jararacussu","volume":"55","author":"Cintra","year":"1991","journal-title":"Exp. Mol. Pathol."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Sanchez-Castro, E.E., Pajuelo-Reyes, C., Tejedo, R., Soria-Juan, B., Tapia-Limonchi, R., Andreu, E., Hitos, A.B., Martin, F., Cahuana, G.M., and Guerra-Duarte, C. (2020). Mesenchymal Stromal Cell-Based Therapies as Promising Treatments for Muscle Regeneration After Snakebite Envenoming. Front. Immunol., 11.","DOI":"10.3389\/fimmu.2020.609961"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"858","DOI":"10.1002\/mus.26329","article-title":"Evidence of induced muscle regeneration persists for years in the mouse","volume":"58","author":"Meyer","year":"2018","journal-title":"Muscle Nerve"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"202","DOI":"10.1002\/mus.23489","article-title":"Homogenates of skeletal muscle injected with snake venom inhibit myogenic differentiation in cell culture","volume":"47","author":"Robledo","year":"2013","journal-title":"Muscle Nerve"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Herrera, C., Mac\u00eado, J.K.A., Feoli, A., Escalante, T., Rucavado, A., Guti\u00e9rrez, J.M., and Fox, J.W. (2016). Muscle Tissue Damage Induced by the Venom of Bothrops asper: Identification of Early and Late Pathological Events through Proteomic Analysis. PLOS Neglected Trop. Dis., 10.","DOI":"10.1371\/journal.pntd.0004599"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1186\/s41232-023-00308-z","article-title":"Extracellular matrix: The critical contributor to skeletal muscle regeneration\u2014A comprehensive review","volume":"43","author":"Ahmad","year":"2023","journal-title":"Inflamm. Regen."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Herrera, C., Escalante, T., Voisin, M.-B., Rucavado, A., Moraz\u00e1n, D., Mac\u00eado, J.K.A., Calvete, J.J., Sanz, L., Nourshargh, S., and Guti\u00e9rrez, J.M. (2015). Tissue Localization and Extracellular Matrix Degradation by PI, PII and PIII Snake Venom Metalloproteinases: Clues on the Mechanisms of Venom-Induced Hemorrhage. PLoS Negl. Trop. Dis., 9.","DOI":"10.1371\/journal.pntd.0003731"},{"key":"ref_47","first-page":"538","article-title":"Morphology of local tissue damage in experimental snake envenomation","volume":"52","author":"Homma","year":"1971","journal-title":"Br. J. Exp. Pathol."},{"key":"ref_48","first-page":"56","article-title":"Primary Carcinoma of the Liver excised by Operation","volume":"16","author":"Wright","year":"1923","journal-title":"Proc. R. Soc. Med."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1747","DOI":"10.1101\/gad.465908","article-title":"Fibrinogen drives dystrophic muscle fibrosis via a TGFbeta\/alternative macrophage activation pathway","volume":"22","author":"Vidal","year":"2008","journal-title":"Genes Dev."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Layfield, H.J., Williams, H.F., Ravishankar, D., Mehmi, A., Sonavane, M., Salim, A., Vaiyapuri, R., Lakshminarayanan, K., Vallance, T.M., and Bicknell, A.B. (2020). Repurposing Cancer Drugs Batimastat and Marimastat to Inhibit the Activity of a Group I Metalloprotease from the Venom of the Western Diamondback Rattlesnake, Crotalus atrox. Toxins, 12.","DOI":"10.3390\/toxins12050309"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Richards, N.J., Alqallaf, A., Mitchell, R.D., Parnell, A., Haidar, H.B., Almeida, J.R., Williams, J., Vijayakumar, P., Balogun, A., and Matsakas, A. (2023). Indian Ornamental Tarantula (Poecilotheria regalis) Venom Affects Myoblast Function and Causes Skeletal Muscle Damage. Cells, 12.","DOI":"10.3390\/cells12162074"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Almeida, J.R., Mendes, B., Pati\u00f1o, R.S.P., Pico, J., Laines, J., Ter\u00e1n, M., Mogoll\u00f3n, N.G.S., Zaruma-Torres, F., Caldeira, C.A.d.S., and da Silva, S.L. (2020). Assessing the stability of historical and desiccated snake venoms from a medically important Ecuadorian collection. Comp. Biochem. Physiol. Part C Toxicol. Pharmacol., 230.","DOI":"10.1016\/j.cbpc.2020.108702"}],"container-title":["Toxins"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-6651\/17\/10\/496\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,15]],"date-time":"2025-10-15T07:39:32Z","timestamp":1760513972000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-6651\/17\/10\/496"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,6]]},"references-count":52,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2025,10]]}},"alternative-id":["toxins17100496"],"URL":"https:\/\/doi.org\/10.3390\/toxins17100496","relation":{},"ISSN":["2072-6651"],"issn-type":[{"value":"2072-6651","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,6]]}}}