LC-MS Analysis and Intracellular and Extracellular Indole 3 Acetic Acid Production under Different Media by Endophytic Bacteria Associated with Humulus lupulus
Main Article Content
Abstract
Recently, there has been a worldwide call to explore nature-friendly metabolites, which could enhance plant growth and substitute for chemically synthesized products. Indole-3-acetic acid (IAA) is one of the versatile metabolites that has a potential role for plant growth, anti-inflammatory, hepatoprotective, and anticancer properties. Furthermore, IAA is commonly synthesized chemically; the majority of reagents used pose environmental pollution. In contrast, biosynthesis through controlled cultivation of endophytes from medicinal plants offers an environmentally sustainable approach. The current study investigates the endophytic bacterium Bacillus licheniformis SKAM1 isolated from the leaves of Humulus lupulus for IAA production. The identification and characterization of endophytic bacterium was carried out using biochemical and molecular methods. Furthermore, LC-MS analysis of the dried extract of Bacillus licheniformis SKAM1 identified multiple bioactive compounds, including IAA, with potential therapeutic and agricultural applications. Additionally, the IAA quantification was performed using ultra-performance liquid chromatography (UPLC) across different media. UPLC analysis reveals that Bacillus licheniformis SKAM1 produces IAA in Luria broth medium; the extracellular IAA concentration was determined to be 1.16 mg/mL, whereas the intracellular level reached 1.11 mg/mL. Similarly, culture in minimal medium with extracellular IAA produced at 0.11 mg/mL and intracellular IAA at 0.06 mg/mL. The current study paves the way for exploring the role of abiotic conditions for cost-effective IAA production.
Article Details

This work is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.
Copyright Transfer Statement
The copyright of this article is transferred to Current Applied Science and Technology journal with effect if and when the article is accepted for publication. The copyright transfer covers the exclusive right to reproduce and distribute the article, including reprints, translations, photographic reproductions, electronic form (offline, online) or any other reproductions of similar nature.
The author warrants that this contribution is original and that he/she has full power to make this grant. The author signs for and accepts responsibility for releasing this material on behalf of any and all co-authors.
Here is the link for download: Copyright transfer form.pdf
References
Allen, M. E., Piefer, A. J., Cole, S. N., Werner, J. J., Benziger, P. T., Grieneisen, L., & Britton, S. J. (2019). Characterization of microbial communities populating the inflorescences of Humulus lupulus L. Journal of the American Society of Brewing Chemists, 77(4), 243-250. https://doi.org/10.1080/03610470.2019.1667739
Altschul, S. F., Gish, W., Miller, W., Myers, E. W., & Lipman, D. J. (1990). Basic local alignment search tool. Journal of Molecular Biology, 215(3), 403-410. https://doi.org/10.1016/S0022-2836(05)80360-2
Alvin, A., Miller, K. I., & Neilan, B. A. (2014). Exploring the potential of endophytes from medicinal plants as sources of antimycobacterial compounds. Microbiological Research, 169(7-8), 483-495. https://doi.org/10.1016/j.micres.2013.12.009
Arora, P., Tabssum, R., Gupta, A. P., Kumar, S., & Gupta, S. (2024). Optimization of indole acetic acid produced by plant growth promoting fungus, aided by response surface methodology. Heliyon, 10(14), Article e34356. https://doi.org/10.1016/j.heliyon.2024.e34356
Atanasov, A. G., Waltenberger, B., Pferschy-Wenzig, E.-M., Linder, T., Wawrosch, C., Uhrin, P., Temml, V., Wang, L., Schwaiger, S., Heiss, E. H., Rollinger, J. M., Schuster, D., Breuss, J. M., Bochkov, V., Mihovilovic, M. D., Kopp, B., Bauer, R., Dirsch, V. M., & Stuppner, H. (2015). Discovery and resupply of pharmacologically active plant-derived natural products: A review. Biotechnology Advances, 33(8), 1582-1614. https://doi.org/10.1016/j.biotechadv.2015.08.001
Basumatary, B., Das, D., Choudhury, B. N., Dutta, P., & Bhattacharyya, A. (2021). Isolation and characterization of endophytic bacteria from tomato foliage and their in vitro efficacy against root-knot nematodes. Journal of Nematology, 53(1), Article e2021-104. https://doi.org/10.21307/jofnem-2021-104
Bocquet, L., Sahpaz, S., & Rivière, C. (2018). An overview of the antimicrobial properties of hop. In J. M. Mérillon and C. Riviere (Eds.). Natural antimicrobial agents. Sustainable development and biodiversity, vol 19 (pp. 31-54). https://doi.org/10.1007/978-3-319-67045-4_2
Bunsangiam, S., Thongpae, N., Limtong, S., & Srisuk, N. (2021). Large scale production of indole-3-acetic acid and evaluation of the inhibitory effect of indole-3-acetic acid on weed growth. Scientific Reports, 11(1), Article 13094. https://doi.org/10.1038/s41598-021-92305-w
Collinge, D. B., Jensen, B., & Jørgensen, H. J. (2022). Fungal endophytes in plants and their relationship to plant disease. Current Opinion in Microbiology, 69, Article 102177. https://doi.org/10.1016/j.mib.2022.102177
de O. Nunes, P. S., de Medeiros, F. H. V., de Oliveira, T. S., de Almeida Zago, J. R., & Bettiol, W. (2023). Bacillus subtilis and Bacillus licheniformis promote tomato growth. Brazilian Journal of Microbiology, 54(1), 397-406. https://doi.org/10.1007/s42770-022-00874-3
Demain, A. L., & Fang, A. (2000). The natural functions of secondary metabolites. In A. Fiechter (Ed.). History of modern biotechnology I. Advances in biochemical engineering/biotechnology. Vol 69 (pp. 1-39). Springer. https://doi.org/10.1007/3-540-44964-7_1
Drlica, K., Zhao, X., & Malik, M. (2009). Quinolones. In Encyclopedia of Microbiology (pp. 707-716). Elsevier. https://doi.org/10.1016/B978-012373944-5.00042-0
Duca, D., Lorv, J., Patten, C. L., Rose, D., & Glick, B. R. (2014). Indole-3-acetic acid in plant–microbe interactions. Antonie van Leeuwenhoek, 106(1), 85-125. https://doi.org/10.1007/s10482-013-0095-y
Emitaro, W. O., Kawaka, F., Musyimi, D. M., & Adienge, A. (2024). Diversity of endophytic bacteria isolated from leguminous agroforestry trees in western Kenya. AMB Express, 14(1), Article 18. https://doi.org/10.1186/s13568-024-01676-6
Folkes, L. K., & Wardman, P. (2001). Oxidative activation of indole-3-acetic acids to cytotoxic species— a potential new role for plant auxins in cancer therapy. Biochemical Pharmacology, 61(2), 129-136. https://doi.org/10.1016/S0006-2952(00)00498-6
Gammoh, O., Aljabali, A. A. A., & Tambuwala, M. M. (2024). Plasma amino acids in major depressive disorder: between pathology to pharmacology. EXCLI Journal, 23, 62-78). https://doi.org/10.17179/excli2023-6767
Goryluk-Salmonowicz, A., Piórek, M., Rekosz-Burlaga, H., Studnicki, M., & Mieczysław, B. (2016). Endophytic detection in selected European herbal plants. Polish Journal of Microbiology, 65(3), 369-375. https://doi.org/10.5604/17331331.1215617
Gouda, S., Das, G., Sen, S. K., Shin, H.-S., & Patra, J. K. (2016). Endophytes: A treasure house of bioactive compounds of medicinal importance. Frontiers in Microbiology, 7, Article 1538. https://doi.org/10.3389/fmicb.2016.01538
Haggag, W. M. (2010). Role of entophytic microorganisms in biocontrol of plant diseases. Life Science Journal, 7(2), 92-97.
Ham, S., Yoon, H., Park, J.-M., & Park, Y. G. (2021). Optimization of fermentation medium for indole acetic acid production by Pseudarthrobacter sp. NIBRBAC000502770. Applied Biochemistry and Biotechnology, 193(8), 2567-2579. https://doi.org/10.1007/s12010-021-03558-0
Hazarika, S. N., Saikia, K., Borah, A., & Thakur, D. (2021). Prospecting endophytic bacteria endowed with plant growth promoting potential isolated from Camellia sinensis. Frontiers in Microbiology, 12, Article 738058. https://doi.org/10.3389/fmicb.2021.738058
Jeong, G.-J., Khan, F., Khan, S., Tabassum, N., Mehta, S., & Kim, Y.-M. (2023). Pseudomonas aeruginosa virulence attenuation by inhibiting siderophore functions. Applied Microbiology and Biotechnology, 107(4), 1019-1038. https://doi.org/10.1007/s00253-022-12347-6
Jeong, G.-J., Khan, S., Tabassum, N., Khan, F., & Kim, Y.-M. (2022). Marine-bioinspired nanoparticles as potential drugs for multiple biological roles. Marine Drugs, 20(8), Article 527. https://doi.org/10.3390/md20080527
Kang, S. M., Shahzad, R., Khan, M. A., Hasnain, Z., Lee, K. E., Park, H. S., Kim, L. R., & Lee, I. J. (2021). Ameliorative effect of indole-3-acetic acid- and siderophore-producing Leclercia adecarboxylata MO1 on cucumber plants under zinc stress. Journal of Plant Interactions, 16(1), 30-41. https://doi.org/10.1080/17429145.2020.1864039
Khan, A. L., Halo, B. A., Elyassi, A., Ali, S., Al-Hosni, K., Hussain, J., Al-Harrasi, A., & Lee, I.-J. (2016). Indole acetic acid and ACC deaminase from endophytic bacteria improves the growth of Solanum lycopersicum. Electronic Journal of Biotechnology, 21, 58-64. https://doi.org/10.1016/j.ejbt.2016.02.001
Khan, N., Ali, S., Shahid, M. A., Mustafa, A., Sayyed, R. Z., & Curá, J. A. (2021). Insights into the interactions among roots, rhizosphere, and rhizobacteria for improving plant growth and tolerance to abiotic Stresses: A review. Cells, 10(6), Article 1551. https://doi.org/10.3390/cells10061551
Khan, S., & Mathur, A. (2025). LC–MS and GC-MS-based bioactive metabolites profiling of endophytic bacterium from Humulus lupulus and production of Indole-3-acetic acid. Current Trends in Biotechnology and Pharmacy, 19(3), 2422-2432. https://doi.org/10.5530/ctbp.2025.3.27
Khan, S., Mathur, A., & Khan, F. (2025). Endophytic fungi-bioinspired nanoparticles potential to control infectious disease. Critical Reviews in Microbiology, 51(6), 1292-1314. https://doi.org/10.1080/1040841X.2025.2497795
Khianngam, S., Meetum, P., Na Chiangmai, P., & Tanasupawat, S. (2023). Identification and optimization of indole-3-acetic acid production of endophytic bacteria and their effects on plant growth. Tropical Life Sciences Research, 34(1), 219-239. https://doi.org/10.21315/tlsr2023.34.1.12
Kholikov, A., Vokhidov, K., Murtozoyev, A., Tóth, Z. S., Nagy, G. N., Vértessy, B. G., & Makhsumkhanov, A. (2025). Characterization of a thermostable α-amylase from Bacillus licheniformis 104.K for industrial applications. Microorganisms, 13(8), Article 1757. https://doi.org/10.3390/microorganisms13081757
Kim, A. Y., Shahzad, R., Kang, S. M., Seo, C. W., Park, Y. G., Park, H. J., & Lee, I. J. (2017). IAA-producing Klebsiella variicola AY13 reprograms soybean growth during flooding stress. Journal of Crop Science and Biotechnology, 20(4), 235-242. https://doi.org/10.1007/s12892-017-0041-0
Kim, J., & Kim, K. H. (2017). Effects of minimal media vs. complex media on the metabolite profiles of Escherichia coli and Saccharomyces cerevisiae. Process Biochemistry, 57, 64-71. https://doi.org/10.1016/j.procbio.2017.04.003
Kitisin, T., Muangkaew, W., Thitipramote, N., Pudgerd, A., & Sukphopetch, P. (2023). The study of tryptophol containing emulgel on fungal reduction and skin irritation. Scientific Reports, 13(1), Article 18881. https://doi.org/10.1038/s41598-023-46121-z
Kolek, J., Patakova, P., Junkova, P., Krofta, K., Hynek, R., & Dostalek, P. (2021). Isolation and identification of Pantoea agglomerans from the inflated bag with dried hop pellets stored under a modified atmosphere. Journal of Applied Microbiology, 131(1), 281-287. https://doi.org/10.1111/jam.14970
Krofta, K., Mravcová, L., Kolek, D., Patáková, P., Patzak, J., Henychová, A., & Dostálek, P. (2021). Microbial community of hop (Humulus lupulus L.) and its impact on the quality of hop products. Acta Horticulturae, 1328, 109-114. https://doi.org/10.17660/ActaHortic.2021.1328.15
Kuźniar, A., Włodarczyk, K., Sadok, I., Staniszewska, M., Woźniak, M., Furtak, K., Grządziel, J., Gałązka, A., Skórzyńska-Polit, E., & Wolińska, A. (2021). A comprehensive analysis using colorimetry, liquid chromatography-tandem mass spectrometry and bioassays for the assessment of indole related compounds produced by endophytes of selected wheat cultivars. Molecules, 26(5), Article 1394. https://doi.org/10.3390/molecules26051394
Lamy, V., Roussi, S., Chaabi, M., Gossé, F., Schall, N., Lobstein, A., & Raul, F. (2007). Chemopreventive effects of lupulone, a hop β-acid, on human colon cancer-derived metastatic SW620 cells and in a rat model of colon carcinogenesis. Carcinogenesis, 28(7), 1575-1581. https://doi.org/10.1093/carcin/bgm080
Li, Y., Li, S., Du, R., Wang, J., Li, H., Xie, D., & Yan, J. (2021). Isoleucine enhances plant resistance against Botrytis cinerea via jasmonate signaling pathway. Frontiers in Plant Science, 12, Article 628328. https://doi.org/10.3389/fpls.2021.628328
Liang, Y., Gong, Y., Yu, Y., Jiang, Q., Zhao, J., Zou, H., & Zhang, J. (2023). Fluorescence resonance energy transfer-based analytical methods for determination of food contaminants: A review. Food Control, 152, Article 109892. https://doi.org/10.1016/j.foodcont.2023.109892
Liu, M., Hansen, P., Wang, G., Qiu, L., Dong, J., Yin, H., Qian, Z., Yang, M., & Miao, J. (2015). Pharmacological profile of xanthohumol, a prenylated flavonoid from hops (Humulus lupulus). Molecules, 20(1), 754-779. https://doi.org/10.3390/molecules20010754
Lu, T., Ke, M., Lavoie, M., Jin, Y., Fan, X., Zhang, Z., Fu, Z., Sun, L., Gillings, M., Peñuelas, J., Qian, H., & Zhu, Y.-G. (2018). Rhizosphere microorganisms can influence the timing of plant flowering. Microbiome, 6(1), Article 231. https://doi.org/10.1186/s40168-018-0615-0
Luziatelli, F., Melini, F., Bonini, P., Melini, V., Cirino, V., & Ruzzi, M. (2021). Production of indole auxins by Enterobacter sp. strain P-36 under submerged conditions. Fermentation, 7(3), Article 138. https://doi.org/10.3390/fermentation7030138
Malinowski, D. P., & Belesky, D. P. (2000). Adaptations of endophyte‐infected cool‐season grasses to environmental stresses: Mechanisms of drought and mineral stress tolerance. Crop Science, 40(4), 923-940. https://doi.org/10.2135/cropsci2000.404923x
Mametja, N. M., Ramadwa, T. E., Managa, M., & Masebe, T. M. (2025). Recent advances and developments in bacterial endophyte identification and application: A 20-Year landscape review. Plants, 14(16), Article 2506. https://doi.org/10.3390/plants14162506
Medison, R. G., Jiang, J., Medison, M. B., Tan, L.-T., Kayange, C. D. M., Sun, Z., & Zhou, Y. (2023). Evaluating the potential of Bacillus licheniformis YZCUO202005 isolated from lichens in maize growth promotion and biocontrol. Heliyon, 9(10), Article e20204. https://doi.org/10.1016/j.heliyon.2023.e20204
Messinis, D. E., Poussin, C., Latino, D. A. R. S., Eb-Levadoux, Y., Dulize, R., Peric, D., Guedj, E., Titz, B., Ivanov, N. V., Peitsch, M. C., & Hoeng, J. (2022). Systems biology reveals anatabine to be an NRF2 activator. Frontiers in Pharmacology, 13, Article 1011184. https://doi.org/10.3389/fphar.2022.1011184
Micci, A., Zhang, Q., Chang, X., Kingsley, K., Park, L., Chiaranunt, P., Strickland, R., Velazquez, F., Lindert, S., Elmore, M., Vines, P. L., Crane, S., Irizarry, I., Kowalski, K. P., Johnston-Monje, D., & White, J. F. (2022). Histochemical evidence for nitrogen-transfer endosymbiosis in non-photosynthetic cells of leaves and inflorescence bracts of angiosperms. Biology, 11(6), Article 876. https://doi.org/10.3390/biology11060876
Mishra, S., Priyanka, & Sharma, S. (2022). Metabolomic insights into endophyte-derived bioactive compounds. Frontiers in Microbiology, 13, Article 835931 https://doi.org/10.3389/fmicb.2022.835931
Mohite, B. (2013). Isolation and characterization of indole acetic acid (IAA) producing bacteria from rhizospheric soil and its effect on plant growth. Journal of Soil Science and Plant Nutrition, 13(3), 638-649. https://doi.org/10.4067/S0718-95162013005000051
Muras, A., Romero, M., Mayer, C., & Otero, A. (2021). Biotechnological applications of Bacillus licheniformis. Critical Reviews in Biotechnology, 41(4), 609-627. https://doi.org/10.1080/07388551.2021.1873239
Narayan, R., Sharma, M., Yadav, R., Biji, A., Khatun, O., Rajmani, R., Sharma, P. R., Jeyasankar, S., Rani, P., Rao, C. D., Satchidanandanam, V., Das, S., Agarwal, R., & Tripathi, S. (2022). A natural broad-spectrum inhibitor of enveloped virus entry, effective against SARS-CoV-2 and influenza A virus in preclinical animal models. https://doi.org/10.1101/2022.02.16.480801
Patakova, P., Vasylkivska, M., Sedlar, K., Jureckova, K., Bezdicek, M., Lovecka, P., Branska, B., Kastanek, P., & Krofta, K. (2024). Whole genome sequencing and characterization of Pantoea agglomerans DBM 3797, endophyte, isolated from fresh hop (Humulus lupulus L.). Frontiers in Microbiology, 15, Article 1305338 https://doi.org/10.3389/fmicb.2024.1305338
Renugadevi, R., Das, M. P. A., Sowndharya, G., Samuel, S. A., Nivethitha, T., Shini, V. S., Sangari, T. L. S., Ramya, M., & Reshma, I. (2022). Production and characterization of indole acetic acid of endophytic bacteria isolated from Kalanchoe pinnata (Lam.) in optimized media. Indian Journal of Science and Technology, 15(38), 1949-1957. https://doi.org/10.17485/IJST/v15i38.1343
Riccioni, C., Belfiori, B., Cenci, M., & Rubini, A. (2025). Exploring endophytic fungi from Humulus lupulus L. for biocontrol of phytopathogenic fungi. Diversity, 17(2), Article 94. https://doi.org/10.3390/d17020094
Rugbjerg, P., Feist, A. M., & Sommer, M. O. A. (2018). Enhanced metabolite productivity of Escherichia coli adapted to glucose M9 minimal medium. Frontiers in Bioengineering and Biotechnology, 6, Article 166. https://doi.org/10.3389/fbioe.2018.00166
Santoyo, G., Moreno-Hagelsieb, G., Orozco-Mosqueda, M. D. C., & Glick, B. R. (2016). Plant growth-promoting bacterial endophytes. Microbiological Research, 183, 92-99. https://doi.org/10.1016/j.micres.2015.11.008
Seo, Y. D., & Wargo, J. A. (2023). From bugs to drugs: Bacterial 3-IAA enhances efficacy of chemotherapy in pancreatic cancer. Cell Reports Medicine, 4(5), Article 101039. https://doi.org/10.1016/j.xcrm.2023.101039
Sevigny, J. L., Lloyd, B., McComish, C., Ramsey, A., & Koziol, L. (2019). Whole-genome sequences of Pantoea agglomerans BL3, Pseudomonas fluorescens BL, and Pseudomonas stutzeri CM14, isolated from hops (Humulus lupulus). Microbiology Resource Announcements, 8(30), Article e00545-19. https://doi.org/10.1128/MRA.00545-19
Sezonov, G., Joseleau-Petit, D., & D’Ari, R. (2007). Escherichia coli Physiology in Luria-Bertani Broth. Journal of Bacteriology, 189(23), 8746-8749. https://doi.org/10.1128/JB.01368-07
Sharma, M., Sood, G., & Chauhan, A. (2021). Bioprospecting beneficial endophytic bacterial communities associated with Rosmarinus officinalis for sustaining plant health and productivity. World Journal of Microbiology and Biotechnology, 37(8), Article 135. https://doi.org/10.1007/s11274-021-03101-7
Shen, J., Yang, L., You, K., Chen, T., Su, Z., Cui, Z., Wang, M., Zhang, W., Liu, B., Zhou, K., & Lu, H. (2022). Indole-3-Acetic acid alters intestinal microbiota and alleviates ankylosing spondylitis in mice. Frontiers in Immunology, 13, Article 762580. https://doi.org/10.3389/fimmu.2022.762580
Shilev, S., Babrikova, I., & Babrikov, T. (2020). Consortium of plant growth‐promoting bacteria improves spinach (Spinacea oleracea L.) growth under heavy metal stress conditions. Journal of Chemical Technology and Biotechnology, 95(4), 932-939. https://doi.org/10.1002/jctb.6077
Shurigin, V., Alimov, J., Davranov, K., Gulyamova, T., & Egamberdieva, D. (2022). The diversity of bacterial endophytes from Iris pseudacorus L. and their plant beneficial traits. Current Research in Microbial Sciences, 3, Article 100133. https://doi.org/10.1016/j.crmicr.2022.100133
Sigurdarson, J. J., Svane, S., & Karring, H. (2020). Development of a M9‐based urea medium (M9U) for sensitive and real‐time monitoring of ureolytic activity of bacteria and cell‐free urease. MicrobiologyOpen, 9(3), article e976. https://doi.org/10.1002/mbo3.976
Singh, M., Kumar, A., Singh, R., & Pandey, K. D. (2017). Endophytic bacteria: a new source of bioactive compounds. 3 Biotech, 7(5), Article 315. https://doi.org/10.1007/s13205-017-0942-z
Singh, R., Thakur, L., Kumar, A., Singh, S., Kumar, S., Kumar, M., Kumar, Y., & Kumar, N. (2023). Comparison of freeze-thaw and sonication cycle-based methods for extracting AMR-associated metabolites from Staphylococcus aureus. Frontiers in Microbiology, 14, Article 1152162. https://doi.org/10.3389/fmicb.2023.1152162
Spaepen, S., Vanderleyden, J., & Remans, R. (2007). Indole-3-acetic acid in microbial and microorganism-plant signaling. FEMS Microbiology Reviews, 31(4), 425-448. https://doi.org/10.1111/j.1574-6976.2007.00072.x
Stofan, M., & Guo, G. L. (2020). Bile acids and FXR: Novel targets for liver diseases. Frontiers in Medicine, 7, Article 544. https://doi.org/10.3389/fmed.2020.00544
Strobel, G., & Daisy, B. (2003). Bioprospecting for microbial endophytes and their natural products. Microbiology and Molecular Biology Reviews, 67(4), 491-502. https://doi.org/10.1128/MMBR.67.4.491-502.2003
Szkop, M., & Bielawski, W. (2013). A simple method for simultaneous RP-HPLC determination of indolic compounds related to bacterial biosynthesis of indole-3-acetic acid. Antonie van Leeuwenhoek, 103(3), 683-691. https://doi.org/10.1007/s10482-012-9838-4
Tao, L., Qiuhong, L., Fuqiang, Y., Shuhui, Z., Suohui, T., & Linyuan, F. (2022). Plant growth-promoting activities of bacterial endophytes isolated from the medicinal plant Pairs polyphylla var. yunnanensis. World Journal of Microbiology and Biotechnology, 38(1), Article 15. https://doi.org/10.1007/s11274-021-03194-0
Teale, W. D., Paponov, I. A., & Palme, K. (2006). Auxin in action: signalling, transport and the control of plant growth and development. Nature Reviews Molecular Cell Biology, 7(11), 847-859. https://doi.org/10.1038/nrm2020
Venkataramanamma, K., Reddy, B. V. B., Jayalakshmi, R. S., Jayalakshmi, V., & Rajendran, L. (2022). Isolation, in vitro evaluation of Bacillus spp. against Fusarium oxysporum f.sp. ciceris and their growth promotion activity. Egyptian Journal of Biological Pest Control, 32(1), Article 123. https://doi.org/10.1186/s41938-022-00618-3
Wary, S., Sarma, A., Talukdar, R., & Tayung, K. (2022). Leaf endophytic fungi of Cymbidium aloifolium L. produces antimicrobials and indole-3-acetic acid. South African Journal of Botany, 149, 381-388. https://doi.org/10.1016/j.sajb.2022.06.035
Weber, S. S., Kaminski, K. P., Perret, J.-L., Leroy, P., Mazurov, A., Peitsch, M. C., Ivanov, N. V., & Hoeng, J. (2019). Antiparasitic properties of leaf extracts derived from selected Nicotiana species and Nicotiana tabacum varieties. Food and Chemical Toxicology, 132, Article 110660. https://doi.org/10.1016/j.fct.2019.110660
Wei, J., Chen, F., Liu, Y., Abudoukerimu, A., Zheng, Q., Zhang, X., Sun, Y., & Yimiti, D. (2020). Comparative metabolomics revealed the potential antitumor characteristics of four endophytic fungi of Brassica rapa L. ACS Omega, 5(11), 5939-5950. https://doi.org/10.1021/acsomega.9b04258
Wu, W., Chen, W., Liu, S., Wu, J., Zhu, Y., Qin, L., & Zhu, B. (2021). Beneficial relationships between endophytic bacteria and medicinal plants. Frontiers in Plant Science, 12, Article 646146. https://doi.org/10.3389/fpls.2021.646146
Zanoli, P., & Zavatti, M. (2008). Pharmacognostic and pharmacological profile of Humulus lupulus L. Journal of Ethnopharmacology, 116(3), 383-396. https://doi.org/10.1016/j.jep.2008.01.011
Zou, L., Wang, Q., Li, M., Wang, S., Ye, K., Dai, W., & Huang, J. (2023a). Culturable bacterial endophytes of Aconitum carmichaelii Debx. were diverse in phylogeny, plant growth promotion, and antifungal potential. Frontiers in Microbiology, 14, Article 1192932. https://doi.org/10.3389/fmicb.2023.1192932
Zou, L., Wang, Q., Wu, R., Zhang, Y., Wu, Q., Li, M., Ye, K., Dai, W., & Huang, J. (2023b). Biocontrol and plant growth promotion potential of endophytic Bacillus subtilis JY-7-2L on Aconitum carmichaelii Debx. Frontiers in Microbiology, 13, Article 1059549. https://doi.org/10.3389/fmicb.2022.1059549