Survival of nematode larvae under exposure to explosion-derived contaminants in war-affected environments
Abstract
Pollution of large areas as a result of military actions in Ukraine has been recorded at least since February 2022, when hazardous toxicants started to be released into environment en mass. Having the ability to bioaccumulate, they concentrate in environmental inanimate-nature objects (soil, natural water bodies). Therefore, the study of how those pollutants behave is of great ecological significance. To determine the effects of toxic releases of explosions on nematode larvae, we used 31 co m pounds at two concentrations (0.1% and 1.0%) and three nematodes ( Strongyloides papillosus , Haemonchus contortus , and Muellerius capillaris ). According to the results, a negative impact on the vitality of the nematode larvae was exerted by 15 toxicants. The greatest nematocidal effects were observed after 24 h exposure to 1% solutions of hydrazine hydrate, 4-nitro- phen ol, p-nitroaniline, and 1-nitro naphthalene, which killed over 80% of the larvae of all three nematodes at different stages. The larvae of the nematodes were also sensitive to 3-nitrophthalic acid dimethyl ester, 1-methyl-2-nitro-benzene, o-nitroaniline, 2-hydroxy-5-nitro-benzaldehyde, 4,5-dibromo-2-nitroaniline, 2-methyl-5-nitro-benzenamine, trinitrotoluene (TNT), (2,4-dinitrophenyl)-h ydrazine, fluorene, N-phenyl-1-naphthalenamine, and pentachloropyridine.References
Ahmad, F., Babalola, O. O., & Siddiqui, M. A. (2012). Integrated approach for management of nematodes in chickpea. Journal of Pure and Applied Microbiology, 6(3), 1063–1068.
Almashat, S., & McDiarmid, M. (2023). Toxic chemical exposures among civilians in armed conflicts: The need for research equity, justice, and accountability. Inhalation Toxicology, 36(5), 304–313.
Bempelou, E., Anagnostopoulos, C., & Liapis, K. (2019). Investigation of naphthalene contamination in olive oil from Greece. Toxicological and Environmental Chemistry, 101, 45–58.
Boiani, M., Merlino, A., Gerpe, A., Porcal, W., Croce, F., Depaula, S., Rodriguez-Haralambides, M. A., Cerecetto, H., & Gonzlez, M. (2009). o-Nitroanilines as major metabolic products of anti-Trypanosoma cruzi 5-phenylethenylbenzofuroxans in microsomal and cytosolic fractions of rat hepatocytes and in whole parasitic cells. Xenobiotica, 39(3), 236–248.
Bomhard, E. M., & Herbold, B. A. (2005). Genotoxic activities of aniline and its metabolites and their relationship to the carcinogenicity of aniline in the spleen of rats. Critical Reviews in Toxicology, 35(10), 783–835.
Boyko, O. O., & Brygadyrenko, V. V. (2019). The viability of Haemonchus contortus (Nematoda, Strongylida) and Strongyloides papillosus (Nematoda, Rhabditida) larvae exposed to various flavourings and source materials used in food production, Vestnik Zoologii, 53(6), 433–442.
Boyko, O. O., & Brygadyrenko, V. V. (2021). Nematicidal activity of aqueous tinctures of plants against larvae of the nematode Strongyloides papillosus. Tropical Biomedicine, 38(2), 85–93.
Boyko, O. O., Gugosyan, Y. A., Shendryk, L. I., & Brygadyrenko, V. V. (2019). Intraspecific morphological variation in free-living stages of Strongyloides papillosus (Nematoda, Strongyloididae) parasitizing various mammal species. Vestnik Zoologii, 53(4), 313–324.
Boyko, O., & Brygadyrenko, V. (2022). Nematicidal activity of organic food additives. Diversity, 14, 615.
Chawla, M. (2025). Environmental contamination and toxicology of benzene-hexachloride (BHC). In: Chawla, M., Singh, J., & Kaushik, R. D. (Eds.). Hazardous chemicals. Overview, toxicological profile, challenges, and future perspectives. Elsevier. Pp. 85–103.
Chen, C., Zhang, Z., Xu, P., Hu, H., & Tang, H. (2023). Anaerobic biodegradation of polycyclic aromatic hydrocarbons. Environmental Research, 223, 115472.
Chen, Q., Li, Z., Chen, Y., Liu, M., Yang, Q., Zhu, B., Mu, J., Feng, L., & Chen, Z. (2024). Effects of electron acceptors and donors on anaerobic biodegradation of PAHs in marine sediments. Marine Pollution Bulletin, 199, 115925.
Chen, T., & Ma, Y. (2025). Enhanced anaerobic biodegradation of PAHs by rhamnolipid and earthworm casts in contaminated soil. Sustainability, 17(12), 5417.
Dave, G. (2000). Sediment and water phase toxicity and UV-activation of six chemicals used in military explosives. Aquatic Ecosystem Health and Management, 3(3), 291–299.
Dhar, K., Abinandan, S., Sana, T., Venkateswarlu, K., & Mallavarapu, M. (2023). Anaerobic biodegradation of phenanthrene and pyrene by sulfate-reducing cultures enriched from contaminated freshwater lake sediments. Environmental Research, 235, 116616.
Dhar, K., Venkateswarlu, K., & Mallavarapu, M. (2024). Enrichment of polycyclic aromatic hydrocarbon (PAH) – degrading strictly anaerobic sulfate-reducing cultures from contaminated soil and sediment. Current Protocols, 4(7), e1102.
Ding, H., Lan, J., Yao, S., Zhang, D., Han, B., Pan, G., & Li, X. (2022). Evolution of polycyclic aromatic hydrocarbons in the surface sediment of Southern Jiaozhou Bay in Northern China after an accident of oil pipeline explosion. Marine Pollution Bulletin, 183, 114039.
Dovhanenko, D. О., Yakovenko, V. M., Brygadyrenko, V. V., & Boyko, O. O. (2024b). Characteristic of the dried-up zone formed as a result of the breach of the Kahovka Dam. Biosystems Diversity, 32(2), 285–295.
Dovhanenko, D. О.,Yakovenko, V. M., Brygadyrenko, V. V., & Boyko, O. O. (2024a). Complex characteristics of landscape components affected by the disaster at the Kahovka Hydropower Plant. Biosystems Diversity, 32(1), 174–182.
Gao, Y., Li, Q., Ling, W., & Zhu, X. (2011). Arbuscular mycorrhizal phytoremediation of soils contaminated with phenanthrene and pyrene. Journal of Hazardous Materials, 185(2–3), 703–709.
Gugosyan, Y. A., Boyko, O. O., & Brygadyrenko, V. V. (2019). Morphological variation of four species of Strongyloides (Nematoda, Rhabditida) parasitising various mammal species. Biosystems Diversity, 27(1), 85–98.
Guo, Z., Kang, Y., Wu, H., Li, M., Hu, Z., & Zhang, J. (2023). Enhanced removal of phenanthrene and nutrients in wetland sediment with metallic biochar: Performance and mechanisms. Chemosphere, 327, 138523.
Heker, I., Samak, N. A., Kong, Y., & Meckenstock, R. U. (2025). Anaerobic degradation of polycyclic aromatic hydrocarbons. Applied and Environmental Microbiology, 91(4), e02268-24.
Higson, F. K. (1992). Microbial degradation of nitroaromatic compounds. Advances in Applied Microbiology, 37, 1–19.
Hill, F. C., Sviatenko, L. K., Gorb, L., Okovytyy, S. I., Blaustein, G. S., & Leszczynski, J. (2012). DFT M06-2X investigation of alkaline hydrolysis of nitroaromatic compounds. Chemosphere, 88(5), 635–643.
Iniaghe, P. O., & Kpomah, E. D. (2023). A comparative analysis on the concentration and potential risk of polycyclic aromatic hydrocarbons in surface water, sediment and soil from a non-crude oil and a crude oil explosion site in the Niger Delta, Nigeria. Chemistry Africa, 6(3), 1633–1653.
Klein, S., Moritz, A., de Carvalho, L., Knaab, T. C., Lungerich, B., Meyer, L., Sandström, J., Thibaud, J. L., de Villiers, K. A., Fidock, D. A., Wittlin, S., Rottmann, M., Held, J., Burckhardt, B. B., & Kurz, T. (2025). Structureactivity relationships of 3hydroxypropanamidines (HPAs) with potent in vivo antimalarial activity. Journal of Medicinal Chemistry, 68(18), 19229–19248.
Knaab, T. C., Moritz, A., de Carvalho, L., Klein, S., Lungerich, B., Lohse, K., Kruse, L., Mombo-Ngoma, G., Orta, L., Thibaud, J. L., de Villiers, K. A., Fidock, D. A., Burckhardt, B. B., Held, J., Wittlin, S., & Kurz, T. (2025). TKK130 is a 3-hydroxy-propanamidine (HPA) with potent antimalarial in vivo activity and a high barrier to resistance. Journal of Medicinal Chemistry, 68(1), 95–107.
Kovalov, O. O., Kolesnyk, Y. M., Sevalniev, A. I., Sharavara, L. P., Hancheva, O. V., Kovalov, K. O., Tyshchenko, T. M., & Tulushev, Y. O. (2025). Peculiarities of the composition of surface and groundwater in Eastern Ukraine during the war: Assessment of environmental and carcinogenic risks. Modern Medical Technology, 17(2), 83–90.
Leffler, P., Brännäs, E., Ragnvaldsson, D., Wingfors, H., & Berglind, R. (2014). Toxicity and accumulation of trinitrotoluene (TNT) and its metabolites in Atlantic salmon alevins exposed to an industrially polluted water. Journal of Toxicology and Environmental Health, Part A, 77(19), 1183–1191.
Li, X., Liu, J., Chen, F., Cheng, Y., Wang, Y., Li, A., Zhai, F., & Sun, Z. (2024). Phytotoxity of polycyclic aromatic hydrocarbons to Salix viminalis L. Pakistan Journal of Botany, 56(2), 703–710.
Lin, D., Chen, Y., Liang, L., Huang, Z., Guo, Y., Cai, P., & Wang, W. (2023). Effects of exposure to the explosive and environmental pollutant 2,4,6-trinitrotoluene on ovarian follicle development in rats. Environmental Science and Pollution Research, 30(42), 96412–96423.
Liu, H., Cheng, L., Hu, Y., Chen, D., Wang, X., Zhang, X., Li, Z., & Wu, Z. (2024). Hepatotoxicity of oral exposure to 2-methyl-4-nitroaniline: Toxicity prediction and in vivo evaluation. Toxicology Letters, 399, 1–8.
Ma, X., & Wu, S. (2024). Oxygenated polycyclic aromatic hydrocarbons in food: Toxicity, occurrence and potential sources. Critical Reviews in Food Science and Nutrition, 64(15), 4882–4903.
Neuwoehner, J., Schofer, A., Erlenkaemper, B., Steinbach, K., Hund-Rinke, K., & Eisentraeger, A. (2007). Toxicological characterization of 2,4,6-trinitrotoluene, its transformation products, and two nitramine explosives. Environmental Toxicology and Chemistry, 26(6), 1090–1099.
Pal, R., Teli, G., Akhtar, M. J., & Purawarga Matada, G. S. P. (2024). Synthetic product-based approach toward potential antileishmanial drug development. European Journal of Medicinal Chemistry, 263, 115927.
Palchykov, V. A., Malaniia, M. M., Tymoshenko, K. I., & Brygadyrenko, V. V. (2025). Organic soil pollutants caused by military activities. Biosystems Diversity, 33(4), e2561.
Palchykov, V., Yakovenko, V., Boyko, O., Maksymenko, M., & Brygadyrenko, V. (2026). Distribution of organic pollutants in Alluvial soils as a result of the destruction of the Kahovka Dam (Ukraine). Environmental Monitoring and Assessment, 198, 676.
Parakhnenko, V. Н., Zadorozhna, О. М., Liakhovska, N. O., & Blahopoluchna, A. H. (2023). Environmental assessment of chemical pollution of soils as a result of the war. Taurian Scientific Herald, 131, 367–373.
Portugal, M. C. S., Altafim, G. L., de Jesus, S. B., Alves, A. V., Valcárcel Rojas, L. A. V., Zanardi-Lamardo, E., Castro, Í. B., Gallucci, F., & Choueri, R. B. (2025). Toxicity of PAHs-enriched sediments on meiobenthic communities under ocean warming and CO2-driven acidification scenarios. Marine Pollution Bulletin, 212, 117489.
Rao, M. A., Di Rauso Simeone, G., Scelza, R., & Conte, P. (2017). Biochar based remediation of water and soil contaminated by phenanthrene and pentachlorophenol. Chemosphere, 186, 193–201.
Reed, L., Büchner, V., & Tchounwou, P. B. (2007). Environmental toxicology and health effects associated with hexachlorobenzene exposure. Reviews on Environmental Health, 22(3), 213.
Rybalka, D. F., & Brygadyrenko, V. V. (2026). Acute toxicity of TNT derivatives and hydrazine-based compounds from explosive and rocket fuel contamination to darkling beetles (Tenebrio molitor and Opatrum sabulosum). PeerJ, 14, e20427.
Salehi-Lisar, S. Y., Deljoo, S., & Harzandi, A. M. (2015). Fluorene and phenanthrene uptake and accumulation by wheat, alfalfa and sunflower from the contaminated soil. International Journal of Phytoremediation, 17(12), 1145–1152.
Shi, Y., Xue, H., Yao, Y., Jing, C., Liu, R., Niu, Q., & Lu, H. (2023). Overcoming methanogenesis barrier to acid inhibition and enhancing PAHs removal by granular biochar during anaerobic digestion. Chemical Engineering Journal, 477, 147229.
Spencer, P. S., & Kisby, G. E. (2021). Role of hydrazine-related chemicals in cancer and neurodegenerative disease. Chemical Research in Toxicology, 34(9), 1953–1969.
Steevens, J. A., Duke, B. M., Lotufo, G. R., & Bridges, T. S. (2002). Toxicity of the explosives 2,4,6-trinitrotoluene, hexahydro-1,3,5-trinitro-1,3,5-triazine, and octahydro-1,3,5,7-tetranitro-1,3,5,7-tetrazocine in sediments to Chironomus tentans and Hyalella azteca: Low-dose hormesis and high-dose mortality. Environmental Toxicology and Chemistry, 21(7), 1475–1482.
Temple, T., Ladyman, M., Mai, N., Galante, E., Ricamora, M., Shirazi, R., & Coulon, F. (2018). Investigation into the environmental fate of the combined insensitive high explosive constituents 2,4-dinitroanisole (DNAN), 1-nitroguanidine (NQ) and nitrotriazolone (NTO) in soil. Science of the Total Environment, 625, 1264–1271.
Van Wyk, J. A., & Mayhew, E. (2013). Morphological identifcation of parasitic nematode infective larvae of small ruminants and cattle: A practical lab guide. Onderstepoort Journal of Veterinary Research, 80(1), 539.
Van Wyk, J., Cabaret, J., & Michael, L. (2004). Morphological identification of nematode larvae of small ruminants and cattle simplified. Veterinary Parasitology, 119(4), 277–306.
Wang, H.-Y., You, J., Yan, Z.-S., Jiang, H.-L., & Ye, H.-X. (2025). Microbial mechanism of magnetite addition on the biodegradation of phenanthrene in sediments. Zhongguo Huanjing Kexue/China Environmental Science, 45(1), 208–222.
Xue, H., Shi, Y., Qiao, J., Li, X., & Liu, R. (2024). Enhancing anaerobic biodegradation of phenanthrene in polluted soil by bioaugmentation and biostimulation: Focus on the distribution of phenanthrene and microbial community analysis. Sustainability, 16(1), 366.
Yang, X., Huan, Z., Zhao, S., & Xi, H. (2023). Study on environmental pollution behavior/fate of ammunition soil and microbial remediation of TNT and its intermediates. Journal of Cleaner Production, 432, 139715.
Zajac, A. M., & Conboy, G. A. (2011). Veterinary clinical parasitology. 8th ed. Willey-Blackwell.
Zhang, Z., Sun, J., Gong, X., Wang, C., & Wang, H. (2023). Anaerobic biodegradation of pyrene and benzo[a]pyrene by a new sulfate-reducing Desulforamulus aquiferis strain DSA. Journal of Hazardous Materials, 459, 132053.
Zhou, N., Guo, H., Zhang, Z., & Wang, H. (2024). The discrepant metabolic pathways of PAHs by facultative anaerobic bacteria under aerobic and nitrate-reducing conditions. Chemosphere, 351, 141230.



