Abstract
Dairy manure composting is widely applied to stabilize organic waste and reduce environmental pollution, yet the behavior of resistance determinants during this process remains insufficiently resolved. In this study, shotgun metagenomic sequencing was used to characterize temporal changes in antibiotic resistance genes (ARGs), metal resistance genes (MRGs), biocide resistance genes (BRGs), mobile genetic elements (MGEs), and microbial community composition during dairy manure composting. Rather than inferring direct mechanistic causation, our analyses focused on identifying statistically supported trends, associations, and co-occurrence patterns across composting stages. We observed a rapid decline in the relative abundance of ARGs compared with MRGs and BRGs during the thermophilic phase, coinciding with increasing temperature, while specific genes such as sul2 persisted throughout the process. Shifts in microbial community composition, particularly changes in the relative dominance of Actinobacteria and Proteobacteria, were significantly associated with variations in resistome profiles. Correlation and network analyses further revealed strong associations among ARGs, MRGs, BRGs, and MGEs, suggesting potential co-selection and horizontal gene transfer linkages without implying direct causal mechanisms. In addition, several opportunistic bacterial genera showed positive associations with aminoglycoside- and macrolide–lincosamide–streptogramin-type ARGs, indicating possible dissemination risks following compost application. Overall, this study provides an integrated, association-based overview of resistome and microbial community dynamics during dairy manure composting and highlights the importance of considering multiple resistance determinants when evaluating composting as a manure management strategy.
Data availability
The datasets used and/or analysed during the current study are available from the corresponding author on reasonable request.
References
WHO. Antimicrobial resistance. World Health Organization. (2020). https://doi.org/10.1016/S0140-6736(20)31898-3
Berendonk, T. U. et al. Tackling antibiotic resistance: The environmental framework. Nat. Rev. Microbiol. 13 (5), 310–317. https://doi.org/10.1038/nrmicro3439 (2015).
Zhu, Y. G. et al. Diverse and abundant antibiotic resistance genes in Chinese swine farms. Proc. Natl. Acad. Sci. U.S.A. 110 (9), 3435–3440. https://doi.org/10.1073/pnas.1222743110 (2013).
Kumar, K., Gupta, S. C., Baidoo, S. K., Chander, Y. & Rosen, C. J. Antibiotic uptake by plants from soil fertilized with animal manure. J. Environ. Qual. 34 (6), 2082–2085. https://doi.org/10.2134/jeq2005.0026 (2005).
He, L. Y. et al. Discharge of swine wastes risks water quality and food safety. Environ. Sci. Technol. 50 (10), 5485–5493. https://doi.org/10.1021/acs.est.5b06035 (2016).
Xie, W. Y., Shen, Q. & Zhao, F. J. Antibiotics and antibiotic resistance from animal manures to soil: A review. Eur. J. Soil. Sci. 69 (1), 181–195. https://doi.org/10.1111/ejss.12494 (2017).
Pal, C., Bengtsson-Palme, J., Rensing, C., Kristiansson, E. & Larsson, D. G. J. BacMet: Antibacterial biocide and metal resistance genes database. Nucleic Acids Res. 42 (D1), D737–D743. https://doi.org/10.1093/nar/gkt1252 (2014).
Arendt, S., Rajic, A., McEwen, S. A. & Reid-Smith, R. J. Antimicrobial use, resistance, and co-selection in food animal production systems. Environ. Int. 180, 108235. https://doi.org/10.1016/j.envint.2024.108235 (2024).
Gillings, M. R. Integrons: Past, present, and future. Microbiol. Mol. Biol. Rev. 78 (2), 257–277. https://doi.org/10.1128/MMBR.00056-13 (2014).
Marti, R. et al. Impact of manure fertilization on the abundance of antibiotic-resistant bacteria and resistance genes in soil and crops. Environ. Sci. Technol. 47 (16), 9172–9180. https://doi.org/10.1021/es401630k (2013).
Zhang, Y. J. et al. Transfer of antibiotic resistance from manure-amended soils to crops. Environ. Sci. Technol. 53 (1), 507–516. https://doi.org/10.1021/acs.est.8b03685 (2019).
Larney, F. J. & Hao, X. A review of composting as a management alternative for beef cattle feedlot manure. J. Environ. Qual. 36 (2), 322–332. https://doi.org/10.2134/jeq2006.0124 (2007).
Li, H., Qiu, Y., Yao, T., Ma, Y. & He, J. Z. Fate of antibiotic resistance genes during composting of livestock manure: A meta-analysis. J. Hazard. Mater. 384, 121315. https://doi.org/10.1016/j.jhazmat.2019.121315 (2020).
Awasthi, M. K. et al. Global trends in composting research: A bibliometric analysis. Bioresour. Technol. 299, 122592. https://doi.org/10.1016/j.biortech.2019.122592 (2020).
Huang, K., Xia, H., Zhang, Y. & Zhu, Y. G. Composting-driven Attenuation of antibiotic resistance genes: Mechanisms and limitations. J. Hazard. Mater. 401, 123355. https://doi.org/10.1016/j.jhazmat.2020.123355 (2021).
Chen, Y., Zhang, H. & Luo, Y. Fate of antibiotic resistance genes during composting of livestock manure. Environ. Pollut. 243, 189–195. https://doi.org/10.1016/j.envpol.2018.08.050 (2018).
Daniel, R., Staley, C. & Sadowsky, M. J. Environmental persistence of metal resistance genes and co-selection with antibiotic resistance. Curr. Opin. Microbiol. 72, 102275. https://doi.org/10.1016/j.mib.2022.102275 (2023).
Li, B. et al. Metagenomic and network analysis reveal wide distribution and co-occurrence of environmental antibiotic resistance genes. ISME J. 9 (11), 2490–2502. https://doi.org/10.1038/ismej.2015.59 (2015).
Zhou, X., Qiu, S. & Tong, C. Thermophilic microbial succession and antibiotic resistance during manure composting. Bioresour. Technol. 344, 126239. https://doi.org/10.1016/j.biortech.2021.126239 (2022).
Ridley, A. M., McCarthy, D. T. & Hamilton, A. J. Thermophilic bacteria and resistance gene persistence in high-temperature treatment systems. Water Res. 250, 120980. https://doi.org/10.1016/j.watres.2023.120980 (2024).
Chen, Y., Li, J., Zhang, Y. & Wang, J. Persistence of sulfonamide resistance genes during manure composting. J. Hazard. Mater. 384, 121433. https://doi.org/10.1016/j.jhazmat.2019.121433 (2020).
Su, J. Q., Yang, Y. & Zhu, Y. G. Selective pressures driving sulfonamide resistance persistence in agricultural environments. Environ. Pollut. 308, 119665. https://doi.org/10.1016/j.envpol.2022.119665 (2022).
Yin, X. et al. ARGs-OAP v2.0 with an expanded SARG database and hidden Markov models for enhancement characterization and quantification of antibiotic resistance genes in environmental metagenomes. Bioinformatics 34 (13), 2263–2270. https://doi.org/10.1093/bioinformatics/bty053 (2018).
Sun, W., Qian, X., Gu, J., Wang, X. J. & Duan, M. L. Metagenomic insights into the fate of antibiotic resistance genes during organic waste composting: Implications for resistome Attenuation and microbial succession. J. Hazard. Mater. 443, 130223. https://doi.org/10.1016/j.jhazmat.2022.130223 (2023).
R Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing. (2020). https://www.r-project.org/
Wickham, H., François, R., Henry, L., Müller, K. & Vaughan, D. dplyr: A grammar of data manipulation [R package]. (2023). https://cran.r-project.org/package=dplyr
Wickham, H. ggplot2: Elegant graphics for data analysis. Springer. (2016). https://doi.org/10.1007/978-3-319-24277-4
Kolde, R. pheatmap: Pretty heatmaps [R package]. (2019). https://cran.r-project.org/package=pheatmap
Huang, H. linkET: Everything is linkable [R package]. (2021). https://github.com/Hy4m/linkET
Pedersen, T. L. ggraph: An implementation of grammar of graphics for graphs and networks [R package]. (2020). https://cran.r-project.org/package=ggraph
Oksanen, J. et al. vegan: Community ecology package [R package]. (2022). https://cran.r-project.org/package=vegan
Qi, L., Li, J. & Zhou, Y. Biocide resistance genes in manure composting systems. Environ. Pollut. 268, 115862. https://doi.org/10.1016/j.envpol.2020.115862 (2021).
Tong, C., Qiu, S. & Wang, Y. Resistome evolution during dairy manure composting. Sci. Total Environ. 807, 150825. https://doi.org/10.1016/j.scitotenv.2021.150825 (2022).
Fu, J., Zhang, T., Li, P. & Wu, D. Thermophilic composting reduces antibiotic resistance genes in livestock manure. Bioresour. Technol. 367, 128266. https://doi.org/10.1016/j.biortech.2022.128266 (2023).
Pan, Y., Liu, R., Zhang, Y. & Wang, H. Temperature-mediated Attenuation of antibiotic resistance genes during composting. J. Environ.Manag. 347, 119046. https://doi.org/10.1016/j.jenvman.2023.119046 (2024).
Tong, C., Xiao, D. & Wang, Y. Effects of composting temperature on antibiotic resistance genes. Chemosphere 224, 757–764. https://doi.org/10.1016/j.chemosphere.2019.02.148 (2019).
Xu, R., Zhang, Y. & Li, J. Fate of resistance genes during composting of animal manure. J. Environ. Sci. 112, 197–206. https://doi.org/10.1016/j.jes.2021.05.029 (2022).
Qiu, S., Tong, C. & Wang, Y. Dynamics of antibiotic resistance genes during thermophilic composting. Bioresour. Technol. 352, 127069. https://doi.org/10.1016/j.biortech.2022.127069 (2022).
Jia, S., Shi, P., Hu, Q. & Li, B. Persistence of sulfonamide resistance genes during composting. Environ. Pollut. 262, 114288. https://doi.org/10.1016/j.envpol.2020.114288 (2020).
Zhang, Y., Li, P. & Wu, D. Selective persistence of sulfonamide resistance during composting. Environ. Res. 216, 114532. https://doi.org/10.1016/j.envres.2022.114532 (2023).
Zhang, M., He, L. & Liu, Y. Co-occurrence patterns of ARGs and MRGs in livestock manure composting. Sci. Total Environ. 711, 134484. https://doi.org/10.1016/j.scitotenv.2019.134484 (2020).
Souza, M. L., Pereira, R. A. & Silva, D. Biocide resistance selection in composting environments. J. Clean. Prod. 349, 131303. https://doi.org/10.1016/j.jclepro.2022.131303 (2022).
Li, B. & Peng, Y. Co-occurrence of antibiotic and metal resistance genes in human-associated pathogens. Environ. Sci. Technol. 51 (9), 5125–5133. https://doi.org/10.1021/acs.est.6b06031 (2017).
Peng, S., Zhang, Y. & Li, X. Co-selection of antibiotic and metal resistance genes in livestock manure composting. Sci. Total Environ. 726, 138639. https://doi.org/10.1016/j.scitotenv.2020.138639 (2020).
Chen, Z. Q. Horizontal gene transfer and environmental dissemination of antibiotic resistance. Environ. Microbiol. Rep. 14 (2), 187–198. https://doi.org/10.1111/1758-2229.13044 (2022).
Peng, S., Chen, L. & Wu, J. Role of mobile genetic elements in resistance gene transfer during composting. Environ. Res. 216, 114691. https://doi.org/10.1016/j.envres.2022.114691 (2023).
Naqvi, S. A., Wang, X. & Li, Y. Mobile genetic elements drive resistance dissemination in agricultural systems. Trends Microbiol. 32 (1), 45–58. https://doi.org/10.1016/j.tim.2023.07.004 (2024).
Shi, X., Hu, H. W. & Zhu, Y. G. Environmental reservoirs and dissemination pathways of resistance genes. Microbiome 11, 98. https://doi.org/10.1186/s40168-023-01491-7 (2023).
Jang, H. M., Yoo, S. & Kim, Y. M. Microbial community dynamics and antibiotic resistance during composting. Waste Manag. 79, 683–691. https://doi.org/10.1016/j.wasman.2018.08.020 (2018).
Matheri, A. N., Belaid, M. & Kumar, S. Thermophilic microbial responses during manure composting. Waste Manag. 154, 246–255. https://doi.org/10.1016/j.wasman.2022.10.019 (2023).
Forsberg, K. J. et al. Bacterial phylogeny structures soil resistomes across habitats. Nature 509 (7502), 612–616. https://doi.org/10.1038/nature13377 (2014).
Walsh, C. Molecular mechanisms that confer antibacterial drug resistance. Nature 406 (6797), 775–781. https://doi.org/10.1038/35021219 (2000).
Su, J. Q., Wei, B., Xu, C. Y., Qiao, M. & Zhu, Y. G. Functional metagenomic characterization of antibiotic resistance genes in agricultural soils. Environ. Sci. Technol. 48 (19), 10987–10995. https://doi.org/10.1021/es502889t (2014).
Wu, D., Zhang, Y. & Luo, Y. Mobile genetic elements as drivers of resistance dissemination in manure. Environ. Pollut. 263, 114461. https://doi.org/10.1016/j.envpol.2020.114461 (2020).
Zhu, Y. G. et al. Diverse and abundant antibiotic resistance genes in agricultural soils. Proc. Natl. Acad. Sci. 118 (8), e2020793118. https://doi.org/10.1073/pnas.2020793118 (2021).
Calero-Cáceres, W., Muniesa, M. & Corominas, L. Fate of antibiotic resistance genes in sewage sludge composting. Sci. Total Environ. 650, 132–140. https://doi.org/10.1016/j.scitotenv.2018.08.361 (2019).
Chen, Y., Li, J., Zhang, Y. & Wang, J. Microbial succession governs antibiotic resistance gene persistence during dairy manure composting. J. Hazard. Mater. 443, 130280. https://doi.org/10.1016/j.jhazmat.2022.130280 (2023).
Fan, X., Li, H., Chen, Q. & Zhu, Y. G. Co-selection of antibiotic and metal resistance during organic waste treatment. Environ. Int. 158, 106919. https://doi.org/10.1016/j.envint.2021.106919 (2022).
Liao, H. et al. Microbial community regulates resistome dynamics during composting. Environ. Sci. Technol. 53 (12), 6824–6834. https://doi.org/10.1021/acs.est.9b00979 (2019).
Chen, Q. L. et al. Long-term field application of sewage sludge increases the abundance of antibiotic resistance genes in soil. Environ. Int. https://doi.org/10.1016/j.envint.2016.03.026 (2016).
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All the authors (Yuan Zhou, Kaiyue Liu, Ping Gong, Jian Wu, Zhuqing Ren*, Erguang Jin) equally contributed to the production of this article.
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Zhou, Y., Liu, K., Gong, P. et al. Integrated metagenomic and 16S rRNA analysis reveals temporal associations between resistance genes and microbial communities during dairy manure composting. Sci Rep (2026). https://doi.org/10.1038/s41598-026-37092-y
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DOI: https://doi.org/10.1038/s41598-026-37092-y