Staphylococcus aureus: Structural biology, virulence determinants, resistance mechanisms, and control strategies
Antibiotic resistance in Staphylococcus aureus
DOI:
https://doi.org/10.62310/liab.v6i2.405Keywords:
Staphylococcus aureus, Antimicrobial resistance, MRSA, Virulence factors, Molecular mechanisms, EpidemiologyAbstract
Staphylococcus aureus is considered one of the most clinically important bacterial pathogens worldwide, and contributes substantially to the burden of infectious diseases in both the human and veterinary populations. The pathogenic activity of S. aureus is mediated by the coordinated action of multiple virulence-associated factors, including toxins, haemolysins, adhesins, immune-evasion proteins and biofilm-forming determinants that facilitate colonisation, tissue invasion, and persistence or chronic infection. Antimicrobial resistance in S. aureus constitutes a substantial challenge, particularly amongst methicillin-resistant (MRSA) strains, in which resistance to β-lactam antibiotics is primarily mediated by the acquisition of the mecA gene, present in the staphylococcal chromosomal cassette mec (SCCmec). This gene encodes an altered penicillin-binding protein (PBP2a), which shows a reduced affinity for β-lactams, allowing for the continued synthesis of peptidoglycan despite exposure to antimicrobials. The widespread dissemination of established MRSA lineages globally highlights the need for coordinated strategies such as the integration of genomic surveillance, the responsible use of antimicrobials, specific infection prevention measures and decolonisation strategies within a One Health framework. The One Health approach recognises the interconnected nature of human, animal, and environmental reservoirs and provides a comprehensive strategy to reduce the emergence and persistence of this pathogen.
References
Aboelnaga N, Elsayed SW, Abdelsalam NA, Salem S, Saif NA, Elsayed M, Ayman S, Nasr M, Elhadidy M. (2024). Deciphering the dynamics of methicillin-resistant Staphylococcus aureus biofilm formation: from molecular signaling to nanotherapeutic advances. Cell Communication and Signaling 22: 188. https://doi.org/10.1186/s12964-024-01511-2
Ahmad-Mansour N, Loubet P, Pouget C, Dunyach-Remy C, Sotto A, Lavigne JP, Molle V. (2021). Staphylococcus aureus toxins: an update on their pathogenic properties and potential treatments. Toxins 13. https://doi.org/10.3390/toxins13100677
Alkuraythi D. (2024). Virulence factors and pathogenicity of Staphylococcus aureus. In: Bustos-Martínez J, Valdez-Alarcón J, Hamdan-Partida A, editors, Advances and Perspectives of Infections Caused by Staphylococcus aureus. IntechOpen. Pp 1–18. https://doi.org/10.5772/intechopen.1006633
Alvarado, Silvia. (2020). Prevalencia de las toxinas PVL, TSST1 y las toxinas exfoliativas en las bacterias de origen comunitario por Staphylococcus aureus y su relación con la evolución clínica de los pacientes, en el periodo entre julio 2018 y junio 2019 en el Hospital San Juan de Dios. Master’s Thesis. Universidad de Costa Rica. https://repositorio.ucr.ac.cr/server/api/core/bitstreams/65f80abe-5bca-41f7-a9ad-abf42b43b57b/content
Alves Carneiro V, Soares Melo R, Mateus Gomes Pereira A, Maria Albuquerque Azevedo Á, Nágila Carneiro Matos M, Mesquita Bastos Cavalcante R, Rhonalty Rocha R, de Queiroz Albuquerque V, Alberto Pérez Guerrero J, Eduardo Aragão Catunda Junior F. (2020). Essential oils as an innovative approach against biofilm of multidrug-resistant Staphylococcus aureus. In: Dincer S, Sumengen M, Arkut A, editors, Bacterial Biofilms. IntechOpen. Pp 1–15. https://doi.org/10.5772/intechopen.91833
Archer NK, Mazaitis MJ, William Costerton J, Leid JG, Powers ME, Shirtliff ME. (2011). Staphylococcus aureus biofilms: properties, regulation and roles in human disease. Virulence 2(5): 445–459. https://doi.org/10.4161/viru.2.5.17724
Aslam M, Raza MN, Hurera A, Iqbal A, Ilyas F, Irum T. (2025). Infection control strategies for preventing methicillin-resistant Staphylococcus aureus-related surgical site infections in Pakistan: a mixed-methods analysis. Cureus Journal of Medical Science 17: e85451. https://doi.org/10.7759/cureus.85451
Bae J, Jin H, Kim J, Park M, Lee J, Kim S. (2021). Molecular characteristics and exotoxins of methicillin-resistant Staphylococcus aureus. Biomedical Science Letters 27(4): 195–207. https://doi.org/10.15616/bsl.2021.27.4.195
Baos Muñoz E. (2017). Caracterización y seguimiento de la resistencia a linezolid en Staphylococcus epidermidis en la unidad de cuidados intensivos del Hospital Clínico San Carlos tras la descripción del primer brote de Staphylococcus aureus linezolid resistente. Doctoral thesis, Universidad Complutense de Madrid, Spain.
Bhattacharya M, Horswill AR. (2024). The role of human extracellular matrix proteins in defining Staphylococcus aureus biofilm infections. FEMS Microbiology Reviews 48(1): 1–18. https://doi.org/10.1093/femsre/fuae002
Bear A, Locke T, Rowland-Jones S, Pecetta S, Bagnoli F, Darton TC. (2023). The immune evasion roles of Staphylococcus aureus protein A and impact on vaccine development. Frontiers in Cellular and Infection Microbiology 13: 1242702. https://doi.org/10.3389/fcimb.2023.1242702
Blanchevoye C, Floquet N, Scandolera A, Baud S, Maurice P, Bocquet O, Blaise S, Ghoneim C, Cantarelli B, Delacoux F, Dauchez M, Efremov RG, Martiny L, Duca L, Debelle L. (2013). Interaction between the elastin peptide VGVAPG and human elastin binding protein. Journal of Biological Chemistry 288(2): 1317–1328. https://doi.org/10.1074/jbc.M112.419929
Blechman SE, Wright ES. (2024). Vancomycin-resistant Staphylococcus aureus (VRSA) can overcome the cost of antibiotic resistance and may threaten vancomycin’s clinical durability. PLoS Pathogens 20: e1012422. https://doi.org/10.1371/journal.ppat.1012422
Boyle-Vavra S, Daum RS. (2007). Community-acquired methicillin-resistant Staphylococcus aureus: the role of Panton-Valentine leukocidin. Laboratory Investigation 87(1): 3–9. https://doi.org/10.1038/labinvest.3700501
Casillas-Ituarte NN, Staats AM, Lower BH, Stoodley P, Lower SK. (2021). Host blood proteins as bridging ligand in bacterial aggregation as well as anchor point for adhesion in the molecular pathogenesis of Staphylococcus aureus infections. Micron 150: 103137. https://doi.org/10.1016/j.micron.2021.103137
Castro Pérez VK, da Costa GM, Sá Guimarães A, Heinemann MB, Pereira Lage A, Seles Dorneles EM. (2020). Relationship between virulence factors and antimicrobial resistance in Staphylococcus aureus from bovine mastitis. Journal of Global Antimicrobial Resistance 22: 792–802. https://doi.org/10.1016/j.jgar.2020.06.010
Chen Y, Sun L, Ba X, Jiang S, Zhuang H, Zhu F, Wang H, Lan P, Shi Q, Wang Z, Chen Y, Shi K, Ji S, Jiang Y, Holmes MA, Yu Y. (2022). Epidemiology, evolution and cryptic susceptibility of methicillin-resistant Staphylococcus aureus in China: a whole-genome-based survey. Clinical Microbiology and Infection 28(1): 85–92. https://doi.org/10.1016/j.cmi.2021.05.024
Chen YJ, Er TK. (2025). Monitoring methicillin-resistant Staphylococcus aureus prevalence in Taiwan: a hospital-based surveillance study from 2022 to 2024. IJID Regions 16: 100702. https://doi.org/10.1016/j.ijregi.2025.100702
Corredor-Rozo ZL, Marquez-Ortiz R, Velandia-Romero ML, Abril D, Madroñero J, Prada LF, Vanegas-Gomez N, García B, Echeverz M, Calderón-Peláez MA, Chaparro-Olaya J, Morales L, Nieto-Clavijo C, Escobar-Perez J. (2025). Deciphering the function of Com_YlbF domain-containing proteins in Staphylococcus aureus. Journal of Bacteriology 207(9): e0006125. https://doi.org/10.1128/jb.00061-25
Cussolim P, Salvi A, Melo A, Melo A. (2021). Mechanisms of resistance of staphylococcus aureus to antibiotics. Revista Facultad Saber 6(12): 831–843.
Di Bella S, Marini B, Stroffolini G, Geremia N, Giacobbe DR, Campanile F, Bartoletti M, Alloisio G, Di Risio L, Viglietti G, Principe L, Costantino V, Busetti M, Zerbato V, Mearelli F, Biolo G, Nunnari A, Cafiero CM, di Masi A. (2025). The virulence toolkit of Staphylococcus aureus: a comprehensive review of toxin diversity, molecular mechanisms, and clinical implications. European Journal of Clinical Microbiology and Infectious Diseases 44(8): 1797–1816. https://doi.org/10.1007/s10096-025-05148-y
Espinosa L, Varela C, Martínez EV, Cano R. (2014). Brotes de enfermedades transmitidas por alimentos. España, 2008–2011 (excluye brotes hídricos). Boletín Epidemiológico Semanal 22(11): 130–145. https://repisalud.isciii.es/rest/api/core/bitstreams/fe6b0f99-70ce-49e7-8bda-781db94a003b/content
Fernandes MG. (2018). Infecciones respiratorias por Staphylococcus aureus: implicación clínica de factores de virulencia y persistencia. Doctoral thesis, Universidad Autónoma de Barcelona, Spain. https://ddd.uab.cat/pub/tesis/2018/hdl_10803_663955/mgf1de1.pdf
Galindo A. (2024). Factores de virulencia de Staphylococcus aureus aislados de encías sanas en alumnos de la FESI. Undergraduate thesis, Universidad Nacional Autónoma de México. https://ru.dgb.unam.mx/server/api/core/bitstreams/2325f701-1699-40fd-bf22-f2f511cd665f/content
Getahun DD, Tarekegn HT, Azene BT, Abebe LT, Belete MA, Tessema TS. (2025). Virulence genes and antibiotic resistance profiling of Staphylococcus species isolated from mastitic dairy cows in and around Bahir Dar, Ethiopia. BMC Microbiology 25: 210. https://doi.org/10.1186/s12866-025-03886-9
Gordon YC, Bae JS, Otto M. (2021). Pathogenicity and virulence of Staphylococcus aureus. Virulence 12: 547–569. https://doi.org/10.1080/21505594.2021.1878688
González-Hernández RA, Valdez-Cruz NA, Trujillo-Roldán MA. (2024). Factors that influence the extraction methods of terpenes from natural sources. Chemical Papers 78(5): 2783–2810. https://doi.org/10.1007/s11696-024-03339-z
Hossain M, Aslan B, Hatoum-Aslan A. (2024). Tandem mobilization of anti-phage defenses alongside SCCmec elements in staphylococci. Nature Communications 15: 8820. https://doi.org/10.1038/s41467-024-53146-z
Idrees M, Sawant S, Karodia N, Rahman A. (2021). Staphylococcus aureus biofilm: morphology, genetics, pathogenesis and treatment strategies. International Journal of Environmental Research and Public Health 18(14): 7602. https://doi.org/10.3390/ijerph18147602
Lehman MK, Nuxoll AS, Yamada KJ, Kielian T, Carson SD, Fey PD. (2019). Protease-mediated growth of Staphylococcus aureus on host proteins is opp3 dependent. mBio 10(2): e02553-18. https://doi.org/10.1128/mBio.02553-18
Linz MS, Mattappallil A, Finkel D, Parker D. (2023). Clinical impact of Staphylococcus aureus skin and soft tissue infections. Antibiotics 12(3): 557. https://doi.org/10.3390/antibiotics12030557
Lucio López D, Proaño Pérez E, Bustillos A. (2026). Estrategias terapéuticas no convencionales frente a microorganismos multirresistentes en ambientes hospitalarios: revisión de literatura. Mediciencias UTA 10(1): 94–106. https://doi.org/10.31243/mdc.uta.v10i1.3100.2026
Maldonado Cabrera BR, Maldonado Cabrera PG. (2024). El desarrollo y los complejos mecanismos de resistencia de S. aureus: una amenaza persistente en la era de los antibióticos. Revista Virtual de la Sociedad Paraguaya de Medicina Interna 11(1): e11132402. https://doi.org/10.18004/rvspmi/2312-3893/2024.e11132402
Méndez Y, Barrera M. (2016). Fisiopatología de la sepsis por gram positivos. Revista Cuarzo 22(1): 51-65. https://doi.org/10.26752/cuarzo.v22.n1.146
Meneses Zurita G. (2022). Descripción fisiopatológica de la descamación cutánea por toxinas exfoliativas estafilocócicas, nuevo enfoque de diagnóstico diferencial. Gaceta Médica Boliviana 45(1): 51–55. https://doi.org/10.47993/gmb.v45i1.365
Mestrovic T, Naghavi M, Robles Aguilar G, Davis Weaver N, Swetschinski LR, Wool EE, Araki DT, Gershberg Hayoon A, Gray B, Han C, Ikuta KS, Rosenblad E, Aalruz H, Abd-Elsalam S, Abedi A, Abolhassani H, Aburuz S, Afzal S, Ahmad A, et al. (2025). The burden of bacterial antimicrobial resistance in the WHO Eastern Mediterranean Region 1990–2021: a cross-country systematic analysis with forecasts to 2050. The Lancet Public Health 10(11): e955–e970. https://doi.org/10.1016/S2468-2667(25)00201-4
Muñoz C, Pinilla G, Navarrete J. (2017). Biopelícula en Staphylococcus spp.: estructura, genética y control. Enfermedades Infecciosas y Microbiología 37(1): 18–29.
Muñoz Cutillas A, Bellón Alonso S, Oujo Álamo E, Rodríguez Tubio-Dapena S, Merino Hernández A, Rodríguez Cimadevilla JL. (2025). Severe sepsis due to Panton-Valentine leukocidin-positive Staphylococcus aureus: case report. Acta Pediátrica de México 46(1): 83–88. https://doi.org/10.18233/apm.v46i1.2719
Naghavi M, Vollset SE, Ikuta KS, Swetschinski LR, Gray AP, Wool EE, Robles Aguilar G, Mestrovic T, Smith G, Han C, Hsu RL, Chalek J, Araki DT, Chung E, Raggi C, Gershberg Hayoon A, Davis Weaver N, Lindstedt PA, Smith AE, et al. (2024). Global burden of bacterial antimicrobial resistance 1990–2021: a systematic analysis with forecasts to 2050. The Lancet 404: 1199–1226. https://doi.org/10.1016/S0140-6736(24)01867-1
Nandhini P, Kumar P, Mickymaray S, Alothaim AS, Somasundaram J, Rajan M. (2022). Recent developments in methicillin-resistant Staphylococcus aureus (MRSA) treatment: a review. Antibiotics 11: 606. https://doi.org/10.3390/antibiotics11050606
Nasser A, Jazireian P, Safari H, Alizadeh-Sani M, Pourmand MR, Azimi T. (2019). Staphylococcus aureus versus neutrophil: scrutiny of ancient combat. Microbial Pathogenesis 131: 259–269. https://doi.org/10.1016/j.micpath.2019.04.026
Nguyen LT, Vogel HJ. (2016). Staphylokinase has distinct modes of interaction with antimicrobial peptides, modulating its plasminogen-activation properties. Scientific Reports 6: 31817. https://doi.org/10.1038/srep31817
Olson ME, Nygaard TK, Ackermann L, Watkins RL, Zurek OW, Pallister KB, Griffith S, Kiedrowski MR, Flack CE, Kavanaugh JS, Kreiswirth BN, Horswill AR, Voyich JM. (2013). Staphylococcus aureus nuclease is an SaeRS-dependent virulence factor. Infection and Immunity 81(4): 1316–1324. https://doi.org/10.1128/IAI.01242-12
Pasachova J, Ramírez S, Munoz L. (2019). Staphylococcus aureus: generalidades, mecanismos de patogenicidad y colonización celular. Nova 17: 25–38. https://doi.org/10.22490/24629448.3631
Patel H, Rawat S. (2023). A genetic regulatory see-saw of biofilm and virulence in MRSA pathogenesis. Frontiers in Microbiology 14: 1204428. https://doi.org/10.3389/fmicb.2023.1204428
Pietrocola G, Nobile G, Rindi S, Speziale P. (2017). Staphylococcus aureus manipulates innate immunity through own and host-expressed proteases. Frontiers in Cellular and Infection Microbiology 7: 166. https://doi.org/10.3389/fcimb.2017.00166
Piewngam P, Otto M. (2024). Staphylococcus aureus colonisation and strategies for decolonisation. The Lancet Microbe 5: e606–e618. https://doi.org/10.1016/S2666-5247(24)00040-5
Popovich KJ, Aureden K, Ham DC, Harris AD, Hessels AJ, Huang SS, Maragakis LL, Milstone AM, Moody J, Yokoe D, Calfee DP. (2023). SHEA/IDSA/APIC practice recommendation: strategies to prevent methicillin-resistant Staphylococcus aureus transmission and infection in acute-care hospitals: 2022 update. Infection Control and Hospital Epidemiology 44: 1039–1067. https://doi.org/10.1017/ice.2023.102
Powers ME, Wardenburg JB. (2014). Igniting the fire: Staphylococcus aureus virulence factors in the pathogenesis of sepsis. PLoS Pathogens 10(2): e1003871. https://doi.org/10.1371/journal.ppat.1003871
Rasmi AH, Ahmed EF, Abdullah Darwish AM, Mahmoud Gad GF. (2022). Virulence genes distributed among Staphylococcus aureus causing wound infections and their correlation to antibiotic resistance. BMC Infectious Diseases 22(1): 652. https://doi.org/10.1186/s12879-022-07624-8
Rasquel-Oliveira FS, Ribeiro JM, Martelossi-Cebinelli G, Costa FB, Nakazato G, Casagrande R, Verri WA. (2025). Staphylococcus aureus in inflammation and pain: update on pathologic mechanisms. Pathogens 14(2): 185. https://doi.org/10.3390/pathogens14020185
Rubio Cieza MY, Romão Bonsaglia EC, Rall dos SVLM, Marcos Veiga, Cirone Silva NC. (2024). Staphylococcal enterotoxins: description and importance in food. Pathogens 13)8): 676. https://doi.org/10.3390/pathogens13080676
Santesmases M. (2021). Microbiología en el siglo XX: infecciones, medicina y biología. In: Viales Hurtado RJ, Rodríguez Sánchez C, editors, Historia de la microbiología en contexto global: estudios de caso de Costa Rica, Argentina y España. Centro de Investigaciones Históricas de América Central (CIHAC). Pp. 1–25. https://digital.csic.es/bitstream/10261/263410/4/Micrbiologia%20en%20el%20siglo%20XX.pdf
Sangucho J, Burgos A, Cruz S, Jaramillo E. (2025). Epidemiology of Methicillin-Resistant Staphylococcus aureus with a “One Health” Approach. Anatomía Digital 8(2): 6–28. https://doi.org/10.33262/anatomiadigital.v8i3.3421
Schwartbeck B, Rumpf CH, Hait RJ, Janssen T, Deiwick S, Schwierzeck V, Mellmann A, Kahl BC. (2024). Various mutations in icaR, the repressor of the icaADBC locus, occur in mucoid Staphylococcus aureus isolates recovered from the airways of people with cystic fibrosis. Microbes and Infection 26(4): 105306. https://doi.org/10.1016/j.micinf.2024.105306
Septimus EJ. (2024). Decolonization to prevent healthcare-associated infections: 2024 state-of-the-art review. Medical Research Archives 12(10): 5782. https://doi.org/10.18103/mra.v12i10.5782
Shao K, Yang Y, Gong X, Chen K, Liao Z, Ojha SC. (2025). Staphylococcal drug resistance: mechanisms, therapies, and nanoparticle interventions. Infection and Drug Resistance 18: 1007–1033. https://doi.org/10.2147/IDR.S510024
Sidel Almache KG, Salas De La Fuente SA, Tapia Santana DC, Novoa Obregon AE, Flores Lozada EP, Rodríguez Revelo MS, Morejon Coello PK, Zapata Paredes PR, Huanca Jumbo KE, Yanez Piedra PA. (2020). Introducción a la microbiología y virología. Mawil, primera edición. Mawil Publicaciones Impresas y Digitales, Quito, Ecuador. https://doi.org/10.26820/978-9942-826-32-9
Speziale P, Pietrocola G. (2020). The multivalent role of fibronectin-binding proteins A and B (FnBPA and FnBPB) of Staphylococcus aureus in host infections. Frontiers in Microbiology 11: 2054. https://doi.org/10.3389/fmicb.2020.02054
Stevens DL, Ma Y, Salmi DB, McIndoo E, Wallace RJ, Bryant AE. (2007). Impact of antibiotics on expression of virulence-associated exotoxin genes in methicillin-sensitive and methicillin-resistant Staphylococcus aureus. The Journal of Infectious Diseases 195: 202–213. https://doi.org/10.1086/510396
Tam K, Torres VJ. (2019). Staphylococcus aureus secreted toxins and extracellular enzymes. Microbiology Spectrum 7(2). https://doi.org/10.1128/microbiolspec.gpp3-0039-2018
Touaitia R, Mairi A, Ibrahim NA, Basher NS, Idres T, Touati A. (2025). Staphylococcus aureus: a review of the pathogenesis and virulence mechanisms. Antibiotics 14(5): 470. https://doi.org/10.3390/antibiotics14050470
Tran PM, Tang SS, Salgado-Pabón W. (2022). Staphylococcus aureus β-toxin exerts anti-angiogenic effects by inhibiting re-endothelialization and neovessel formation. Frontiers in Microbiology 13: 840236. https://doi.org/10.3389/fmicb.2022.840236
Truong-Bolduc QC, Yonker LM, Wang Y, Lawton BG, Hooper DC. (2024). NorA efflux pump mediates Staphylococcus aureus response to Pseudomonas aeruginosa pyocyanin toxicity. Antimicrobial Agents and Chemotherapy 68(2): e01001-23. https://doi.org/10.1128/aac.01001-23
Vandenesch F, Lina G, Henry T. (2012). Staphylococcus aureus hemolysins, bi-component leukocidins, and cytolytic peptides: a redundant arsenal of membrane-damaging virulence factors? Frontiers in Cellular and Infection Microbiology 2: 12. https://doi.org/10.3389/fcimb.2012.00012
Visan AI, Negut I. (2024). Coatings based on essential oils for combating antibiotic resistance. Antibiotics 13(7): 625. https://doi.org/10.3390/antibiotics13070625
Visansirikul S, Kolodziej SA, Demchenko AV. (2020). Staphylococcus aureus capsular polysaccharides: a structural and synthetic perspective. Organic and Biomolecular Chemistry 18(5): 783–798. https://doi.org/10.1039/c9ob02546d
Wang J, Li H, Pan J, Dong J, Zhou X, Niu X, Deng X. (2018). Oligopeptide targeting sortase A as potential anti-infective therapy for Staphylococcus aureus. Frontiers in Microbiology 9: 245. https://doi.org/10.3389/fmicb.2018.00245
Xu H, Wang S, Liu X, Li M, Wang X, Chen H, Qu C, Liu Y, Liu J. (2025). Strategies for survival of Staphylococcus aureus in host cells. International Journal of Molecular Sciences 26(2): 720. https://doi.org/10.3390/ijms26020720
Yuan H, Xu J, Wang Y, Li Y, Hao Y, Long J, Liu F, Zhu J, Yang H. (2025). The global antimicrobial resistance trends of Staphylococcus aureus and influencing factors. Microbiology Research 16(6): 118. https://doi.org/10.3390/microbiolres16060118
Zheng M, Zhu K, Peng H, Shang W, Zhao Y, Lu S, Rao X, Li M, Zhou R, Li G. (2022). CcpA regulates Staphylococcus aureus biofilm formation through direct repression of staphylokinase expression. Antibiotics 11(10): 1426. https://doi.org/10.3390/antibiotics11101426
Zumba Camila Berenice R, Bravo Paola Patricia O, Tacuri Carlos Fernando A, Dávila María del Cisne C. (2022). Detection of Staphylococcus aureus in high-speed dental handpieces. Revista OACTIVA UC Cuenca 7(2): 21–27. https://doi.org/10.31984/oactiva.v7i2.746
Zurita F, Hernández M, Romero J, Salazar J, Álvarez R, Galarza G. (2022). Microorganismos y factores relacionados a las infecciones adquiridas en un hospital docente durante los años 2014 a 2018. Revista Médica Vozandes 33: 35–43. https://revistamedicavozandes.com/wp-content/uploads/2023/01/02_AO4.pdf
Downloads
Published
How to Cite
Issue
Section
License
Copyright (c) 2026 Isabel Cristina López Villacís, Sandra Margarita Cruz-Quintana, Jenny Piedad Lozada-Ortiz, Blanca Jeaneth Villavicencio-Villavicencio, Pedro Díaz-Sjostrom, Carlos Vásquez, Andrea Carolina Vela-Chiriboga, Ana Rafaela Burgos Mayorga
This work is licensed under a Creative Commons Attribution 4.0 International License.
Accepted 2026-10-11
Published 2026-10-25
