MMSL X:X | DOI: 10.31482/mmsl.2026.001

BIOLOGICAL WEAPONS – HISTORY AND PRESENTReview article

Macela Aleš ORCID...1, Boštíková Vanda ORCID...2, Kubelková Klára ORCID...3*
1 Military Faculty of Medicine, University of Defense, Třebešská 1575, 500 01 Hradec Králové, Czech Republic
2 Department of Epidemiology, Military Faculty of Medicine, University of Defense, Třebešská 1575, 500 01 Hradec Králové, Czech Republic
3 Department of Molecular Pathology and Biology, Military Faculty of Medicine, University of Defense, Třebešská 1575, 500 01 Hradec Králové, Czech Republic

In 2025, we commemorated the 50th anniversary of the ratification of the Convention on the Prohibition of the Development, Production and Stockpiling of Bacteriological (Biological) and Toxin Weapons and on their Destruction (Biological Weapons Convention, BWC). By 2025, 189 countries had signed the treaty, including the four original signatory states. However, research on microorganisms or their products – toxins – that can be misused for military or terrorist purposes is still being carried out in military facilities in many countries. For this reason, constant attention should be paid to the issues related to the individual Articles of the convention. This article, taking advantage of this anniversary, seeks to provide a precise definition of biological weapons, a brief historical overview of the use of biological agents in armed conflict, and a brief description of the efforts to prohibit the use of microorganisms or their products as weapons of mass destruction, especially against human targets. In its concluding remarks, given the overlap between biomedical and military research on the use of biological agents. The text addresses the still not completely resolved issue of the misuse of biological agents, referring to the long and layered history of their use for nefarious purposes, contributes to distinguish between defensive and offensive biological research, and draws attention to the complexity of issues related to the biological security of world´s states.

Keywords: Biological Weapons Convention; biological agents; biological warfare; history

Received: December 15, 2025; Revised: April 16, 2026; Accepted: April 16, 2026; Prepublished online: April 30, 2026 

Download citation

References

  1. Convention on the Prohibition of the Development, Production and Stockpiling of Bacteriological (Biological) and Toxin Weapons and on Their Destruction, 1972. https://disarmament.unoda.org/biological-weapons/
  2. Millett PD. The biological and toxin weapons convention. Rev Sci Tech Int Off Epizoot. 2006;25(1):35-52.
  3. Ouagrham-Gormley SB. Safeguarding against biological weapons. Science. 2025;388(6749):797. Go to original source... Go to PubMed...
  4. Sundelson AE, Gronvall GK, Ackerman G, et al. Diplomacy disrupted: A mixed-methods analysis of Russian disinformation at the Ninth Review Conference of the Biological and Toxin Weapons Convention. Polit Life Sci. 2025;44(1):28-48. Go to original source... Go to PubMed...
  5. Cavaillon JM. From Bacterial Poisons to Toxins: The Early Works of Pasteurians. Toxins. 2022;14(11):759. Go to original source... Go to PubMed...
  6. Demain AL, Elander RP. The beta-lactam antibiotics: past, present, and future. Antonie Van Leeuwenhoek. 1999;75(1-2):5-19. Go to original source...
  7. Vanĕk Z, Mikulík K. 1978. Microbial growth and production of antibiotics. Folia Microbiol (Prague). 1978;23(4):309-328. Go to original source...
  8. Doehlemann G, Ökmen B, Zhu W, et al. Plant pathogenic fungi. Microbiology spectrum. 2017;5(1). Article FUNK-0023-2016. https://doi.org/10.1128/microbiolspec.FUNK-0023-2016 Go to original source... Go to PubMed...
  9. Ahmad-Mansour N, Loubet P, Pouget C, et al. Staphylococcus aureus Toxins: An Update on Their Pathogenic Properties and Potential Treatments. Toxins. 2021;13(10):677. Go to original source... Go to PubMed...
  10. Wang X, Yang J, Huang P, et al. Cytisine: State of the art in pharmacological activities and pharmacokinetics. Biomed Pharmacother. 2024;171:116210. Go to original source... Go to PubMed...
  11. Hotti H, Rischer H. The killer of Socrates: Coniine and Related Alkaloids in the Plant Kingdom. Molecules. 2017;22(11):1962. doi: 10.3390/molecules22111962. PMID: 29135964; PMCID: PMC6150177. Go to original source...
  12. Xu S, Gaquerel E. Evolution of plant specialized metabolites: beyond ecological drivers. Trends Plant Sci. 2025;30(8):826-836. Go to original source... Go to PubMed...
  13. Giovannini M, Mori F, Barni S, et al. Hymenoptera venom allergy in children. Ital J Pediatr. 2024;50(1):262. Go to original source...
  14. Dos Santos Arraes DR, Rodrigues ABL, Sanches PR, et al. Bioactive alkaloids from the venom of Dendrobatoidea Cope, 1865: a scoping review. J Toxicol Environ Health B Crit Rev. 2024;27(1):1-20. Go to original source... Go to PubMed...
  15. Biological weapons. https://www.who.int/health-topics/biological-weapons#tab=tab_1.
  16. Barras V, Greub G. History of biological warfare and bioterrorism. Clin Microbiol Infect. 2014;20(6):497-502. doi: 10.1111/1469-0691.12706. PMID: 24894605. Go to original source...
  17. Trevisatano S. The 'Hittite plaque' an epidemic of tularemia and the first record of biological warfare. Med Hypotheses. 2017;(69):1371-1374. Go to original source... Go to PubMed...
  18. Conrad LI. The Biblical Tradition for the Plague of the Philistines. J Am Orient Soc. 1984;104(2):281. Go to original source...
  19. Grmek M. Les ruses de guerre biologiques dans l'antiquité. Rev Etud Grec. 1979;92:142-163. French. PMID: 11632888. Go to original source...
  20. Das S, Kataria VK. Bioterrorism: A Public Health Perspective. Med J Armed Forces India. 2010;66(3):255-260. Go to original source... Go to PubMed...
  21. https://greekreporter.com/2025/06/24/mass-poisoning-kirrha-ancient-greece/.
  22. Winter C. War-Canoes and Poisoned Arrows: Great Jolof and Imperial Mali Against the Fifteenth Century Portuguese Slave Raids. Journal of African Military History. 2023;8(1),27-57. https://doi.org/10.1163/24680966-bja10016 Go to original source...
  23. History of Sierra Leone. Embassy of Sierra Leone, Saudi Arabia. Available from: https://sa.slembassy.gov.sl/history-of-sierra-leone/
  24. Jokinen A. 2009. Sir John Hawkyns (1532-1596). Luminarium:Encyklopedia project. Available from: https://luminarium.org/encyclopedia/hawkyns.htm
  25. Papagrigorakis MJ, Synodinos PN, Stathi A, et al. 2013.The plague of Athens: an ancient act of bioterrorism? Biosecurity Bioterrorism Biodefense Strategy Pract Sci. 2013;11(3):228-229. Go to original source... Go to PubMed...
  26. Poole JC, Holladay AJ. Thucydides and the plague of Athens. Class Q. 1979;29:282-300. Go to original source...
  27. McSherry J, Kilpatrick R. The plague of Athens. J R Soc Med. 1992;85(11):713.
  28. Théodoridès J. The plague of Athens. J R Soc Med. 1993;86(4):244.
  29. Theodorides J. The plague of Athens. J R Soc Med. 1995;88(6):363.
  30. Domingo P, Prieto P, Pons L. A plague like no other: beyond the buboes in Thucydides' account of the Plague of Athens Clinical Microbiology and Infection, 2025;31,1639-1646. Go to original source... Go to PubMed...
  31. https://www.historyskills.com/classroom/ancient-history/hannibal-snake-bombs/
  32. Burton A. Bithinian snake bombs. Front Ecol Environ. 2021;19(3):196-197. Go to original source...
  33. https://www.scribd.com/document/321408134/Tactica-of-Emperor-Leo-VI-the-Wise
  34. https://www.thunleveque.fr/notre-commune/historique-de-thun-leveque/
  35. Wheelis M. Biological Warfare at the 1346 Siege of Caffa. Emerg Infect Dis. 2021;8(9):971-975. Go to original source... Go to PubMed...
  36. Derbes VJ. The great plague of 1348. A forgotten episode of bacteriological warfare. JAMA. 1966;196(1):59-62. Go to original source...
  37. Norris J. East or West? The Geographic Origin of the Black Death. Bull Hist Med. 1977;51(1):1-24.
  38. Motyčka J. The royal missions of Sigismund Korybut in Hussite Bohemia and Moravia in a broader geopolitical context. Husité, 2005, (Czech), on-line: https://husitstvi.cz/clanky/vladarske-mise-zikmunda-korybuta-v-husitskych-cechach-a-na-morave-v-sirsich-geopolitickych-souvislostech/
  39. Riedel S. Biological warfare and bioterrorism: a historical review. Proc Bayl Univ Med Cent. 2004;17(4):400-406. Go to original source... Go to PubMed...
  40. Poupard JA, Miller LA. History of biological warfare: catapults to capsomeres. Ann N Y Acad Sci. 1992;31;666:9-20. Go to original source... Go to PubMed...
  41. Carus WS. The History of Biological Weapons Use: What We Know and What We Don't. Health Secur. 2015;13(4):219-255. Go to original source...
  42. Christopher GW, Cieslak TJ, Pavlin JA, et al. Biological warfare. A historical perspective. JAMA. 1997;278(5):412-417. Go to original source...
  43. Society M. Typhus in World War I [Internet]. Available from: https://microbiologysociety.org/publication/past-issues/world-war-i/article/typhus-in-world-war-i.html
  44. Biological and toxin weapons research, development and use from the Middle Ages to 1945. Available from: https://www.google.com/search?client=firefox-b-d&q=Biological+and+toxin+weapons+research%2C+development+and+use+from+the+middle+ages+to+1945
  45. Gruinard Island - an overview | ScienceDirect Topics . Available from: https://www.sciencedirect.com/topics/medicine-and-dentistry/gruinard-island
  46. The Burry Inlet Anthrax Bomb Test 1942. Available from: https://www.llanellich.org.uk/files/319-anthrax
  47. Geißler E. Alibek, Tularaemia and The Battle of Stalingrad. CBW Conv Bull. 2005;1(3),56-68.
  48. Leitenberg M, Zilinskas RA, Kuhn, JH. 2012. The Soviet Biological Weapons Program: A History. Harvard University Press. https://doi.org/10.2307/j.ctt2jbscf Go to original source...
  49. History of three mobilizations: a reexamination of the Chinese biological warfare allegations against the United States in the Korean War - Search Google. Available from: https://www.google.com/search?client=firefox-b-d&q=History+of+three+mobilizations%3A+a+reexamination+of+the+Chinese+biological+warfare+allegations+against+the+United+States+in+the+Korean+War
  50. Deceiving the deceivers: Moscow, Beijing, Pyongyang, and the allegations of bacteriological weapons use in Korea. - Search Google. Available from: https://www.google.com/search?client=firefox-b-d&q=Deceiving+the+deceivers%3A+Moscow%2C+Beijing%2C+Pyongyang%2C+and+the+allegations+of+bacteriological+weapons+use+in+Korea.+
  51. Alibek K, Handelman S. Biohazard. 2000. The Chilling True Story of the Largest Covert Biological Weapons Program in the World--Told from Inside by the Man Who Ran It. New York, NY: Delta; 336.
  52. Domaradskij IV, Orent W. 2003. Biowarrior: Inside the Soviet/Russian Biological War Machine. Amherst, NY: Prometheus;250.
  53. The Nixon administration's decision to end U.S. biological warfare programs. - Search Google. Available from: https://www.google.com/search?client=firefox-b-d≻a_esv=788002144cbb21b9&q=The+Nixon+administration%27s+decision+to+end+U.S.+biological+warfare+programs.&spell=1&sa=X&ved=2ahUKEwit8fLFnrWPAxWIzwIHHcwiH5wQBSgAegQIHxAB&biw=1869&bih=921&dpr=1
  54. Christopher GW, Cieslak TJ, Pavlin JA, et al. Biological warfare. A historical perspective. JAMA. 1997;278(5):412-417. Go to original source...
  55. https://sujb.gov.cz/zakaz-biologickych-zbrani/umluva-o-zakazu-vyvoje-vyroby-a-hromadeni-zasob-bakteriologickych-biologickych-a-toxinovych-zbrani-a-o-jejich-zniceni.
  56. Harris ED. Sverdlovsk and Yellow Rain: Two Cases of Soviet Noncompliance? Int Secur. 1987;11(4):41-95. Go to original source...
  57. Macela A. 2008. Current and Future Developments for Establishing an Operational BIO DIM Capability. BIODIM, Biological detection, identification and monitoring. Brochure EDA symposium, 9-11 June, 2008, Munster, Germany: 4
  58. Biological weapons and potential indicators of offensive biological weapon activities. - Search Google. Available from: https://www.google.com/search?client=firefox-b-d&q=Biological+weapons+and+potential+indicators+of+offensive+biological+weapon+activities.+
  59. Leitenberg M. Distinguishing offensive from defensive biological weapons research. Crit Rev Microbiol. 2003;29(3):223-257. doi: 10.1080/713610450. PMID: 14582619. Go to original source...