Genetic Tolerance of Peanut (Arachis hypogaea L.) to Diverse Aspergillus flavus Strains and Its Implications for Fungal Colonization and Seed Quality in the Drylands of West Nusa Tenggara
DOI:
10.29303/josdas.v6i1.1712Published:
2026-06-30Downloads
Abstract
Peanut (Arachis hypogaea L.) cultivated in tropical dryland agroecosystems faces a serious threat from Aspergillus flavus infection, which produces carcinogenic aflatoxins. This study aimed to synthesize host genetic tolerance mechanisms, pathogen strain diversity, and their interaction affecting fungal colonization and seed quality under dryland stress. A narrative review approach was applied to seventeen primary articles and seven supporting studies selected from PubMed, ProQuest, and ScienceDirect by adopting selected reporting elements of PRISMA 2020, then synthesized through four thematic clusters. The synthesis revealed that peanut tolerance is hierarchical and complementary across three defense levels, namely physical defense based on testa architecture, biochemical defense based on flavonoid accumulation (44.29% of total metabolites) and phenolic acids, and enzymatic-molecular defense through expression of eighteen hub genes and the candidate gene AhAftr1. Strain diversity of A. flavus at the morphotype, population structure, and pangenome levels (59% accessory genes) was closely associated with virulence and aflatoxin production variation. A study under semi-arid conditions reported that drought stress increased aflatoxin B1 contamination by up to 54.95%, with the pod-filling stage as the critical period. An integrated layered control strategy combining multi-trait breeding, atoxigenic biocontrol, and agronomic management is proposed as a site-specific direction for the drylands of West Nusa Tenggara, particularly Lombok Island, and still requires field validation
Keywords:
Aflatoxin Dryland Genetic tolerance Plant breeding Seed coat polyphenolReferences
Ajmal, M., Bedale, W., Akram, A., & Yu, J. H. (2022). Comprehensive Review of Aflatoxin Contamination, Impact on Health and Food Security, and Management Strategies in Pakistan. In Toxins (Vol. 14, Number 12). MDPI. https://doi.org/10.3390/toxins14120845
Akullo, J. O., Okello, D. K., Mohammed, A., Muyinda, R., Amayo, R., Magumba, D., Gidoi, R., Njoroge, S., & Mweetwa, A. (2025). A Comprehensive Review of Aflatoxin in Groundnut and Maize Products in Africa: Prevalence, Detection and Mitigation Strategies. In Journal of Food Quality (Vol. 2025, Number 1). John Wiley and Sons Inc. https://doi.org/10.1155/jfq/2810946
Al-Zaban, M. I. (2023). Impacts of Temperature and Water Activity Interactions on Growth, Aflatoxin B1 Production and Expression of Major Biosynthetic Genes of AFB1 in Aspergillus flavus Isolates. Microorganisms, 11(5). https://doi.org/10.3390/microorganisms11051199
Aminou, M. M., Falalou, H., Abdou, H., & Mendu, V. (2024). Aflatoxin B1 Contamination Association with the Seed Coat Biochemical Marker Polyphenol in Peanuts Under Intermittent Drought. Journal of Fungi, 10(12). https://doi.org/10.3390/jof10120850
Badan Pangan Nasional. (2024). Buku Statistik Ketahanan Pangan 2024. https://data.badanpangan.go.id/statisticpublications/pzg
BMKG. (2024). Climate Outlook 2024. https://www.bmkg.go.id/iklim/climate-outlook-2024
Braun, V., & Clarke, V. (2024). Supporting best practice in reflexive thematic analysis reporting in Palliative Medicine: A review of published research and introduction to the Reflexive Thematic Analysis Reporting Guidelines (RTARG). Palliative Medicine, 38(6), 608–616. https://doi.org/10.1177/02692163241234800
Broto, W. (2018). Status Cemaran Dan Upaya Pengendalian Aflatoksin Pada Komoditas Serealia Dan Aneka Kacang. Jurnal Penelitian Dan Pengembangan Pertanian, 37(2), 81. https://doi.org/10.21082/jp3.v37n2.2018.p81-90
Chai, P., Cui, M., Zhao, Q., Chen, L., Guo, T., Guo, J., Wu, C., Du, P., Liu, H., Xu, J., Zheng, Z., Huang, B., Dong, W., Han, S., & Zhang, X. (2024). Genome-Wide Characterization of the Phenylalanine Ammonia-Lyase Gene Family and Their Potential Roles in Response to Aspergillus flavus L. Infection in Cultivated Peanut (Arachis hypogaea L.). Genes, 15(3). https://doi.org/10.3390/genes15030265
Commey, L., Tengey, T. K., Cobos, C. J., Dampanaboina, L., Dhillon, K. K., Pandey, M. K., Sudini, H. K., Falalou, H., Varshney, R. K., Burow, M. D., & Mendu, V. (2021). Peanut seed coat acts as a physical and biochemical barrier against aspergillus flavus infection. Journal of Fungi, 7(12). https://doi.org/10.3390/jof7121000
Cuartero, J., Querejeta, J. I., Prieto, I., Frey, B., & Alguacil, M. M. (2024). Warming and rainfall reduction alter soil microbial diversity and co-occurrence networks and enhance pathogenic fungi in dryland soils. Science of the Total Environment, 949. https://doi.org/10.1016/j.scitotenv.2024.175006
Cui, M., Han, S., Wang, D., Haider, M. S., Guo, J., Zhao, Q., Du, P., Sun, Z., Qi, F., Zheng, Z., Huang, B., Dong, W., Li, P., & Zhang, X. (2022). Gene Co-expression Network Analysis of the Comparative Transcriptome Identifies Hub Genes Associated With Resistance to Aspergillus flavus L. in Cultivated Peanut (Arachis hypogaea L.). Frontiers in Plant Science, 13. https://doi.org/10.3389/fpls.2022.899177
Dinas Pertanian dan Perkebunan. (2023). Dataset - Satu Data NTB. https://data.ntbprov.go.id/dataset/9d37d7cc-8d48-48cf-b222-ee6d95be6a9a/show
Drott, M. T., Satterlee, T. R., Skerker, J. M., Pfannenstiel, B. T., Glass, N. L., Keller, N. P., & Milgroom, M. G. (2020). The Frequency of Sex: Population Genomics Reveals Differences in Recombination and Population Structure of the Aflatoxin-Producing Fungus Aspergillus flavus. https://doi.org/10.1128/mBio
Gangurde, S. S., Korani, W., Bajaj, P., Wang, H., Fountain, J. C., Agarwal, G., Pandey, M. K., Abbas, H. K., Chang, P. K., Holbrook, C. C., Kemerait, R. C., Varshney, R. K., Dutta, B., Clevenger, J. P., & Guo, B. (2024). Aspergillus flavus pangenome (AflaPan) uncovers novel aflatoxin and secondary metabolite associated gene clusters. BMC Plant Biology, 24(1). https://doi.org/10.1186/s12870-024-04950-8
Gilbert, M. K., Mack, B. M., Moore, G. G., Downey, D. L., Lebar, M. D., Joardar, V., Losada, L., Yu, J. J., Nierman, W. C., & Bhatnagar, D. (2018). Whole genome comparison of aspergillus flavus L-morphotype strain NRRL 3357 (type) and S-morphotype strain AF70. PLoS ONE, 13(7). https://doi.org/10.1371/journal.pone.0199169
Jayaprakash, A., Roy, A., Thanmalagan, R. R., Arunachalam, A., & PTV, L. (2021). Immune response gene coexpression network analysis of Arachis hypogaea infected with Aspergillus flavus. Genomics, 113(5), 2977–2988. https://doi.org/10.1016/j.ygeno.2021.06.027
Jin, G., Liu, N., Yu, B., Jiang, Y., Luo, H., Huang, L., Zhou, X., Yan, L., Kang, Y., Huai, D., Ding, Y., Chen, Y., Wang, X., Jiang, H., Lei, Y., Shen, J., & Liao, B. (2023). Identification and Pyramiding Major QTL Loci for Simultaneously Enhancing Aflatoxin Resistance and Yield Components in Peanut. Genes, 14(3). https://doi.org/10.3390/genes14030625
Joshi, P., Sharma, V., Pandey, A. K., Nayak, S. N., Bajaj, P., Sudini, H. K., Sharma, S., Varshney, R. K., & Pandey, M. K. (2025). Identification of miRNAs associated with Aspergillus flavus infection and their targets in groundnut (Arachis hypogaea L.). BMC Plant Biology 2025 25:1, 25(1), 345-. https://doi.org/10.1186/S12870-025-06322-2
Khadgi, A., Lekkala, S., Verma, P. K., Puppala, N., & Janga, M. R. (2025). Emerging Strategies for Aflatoxin Resistance in Peanuts via Precision Breeding. In Toxins (Vol. 17, Number 8). Multidisciplinary Digital Publishing Institute (MDPI). https://doi.org/10.3390/toxins17080394
Mamo, F. T., Shang, B., Selvaraj, J. N., Zheng, Y., & Liu, Y. (2022). Biocontrol efficacy of atoxigenic Aspergillus flavus strains against aflatoxin contamination in peanut field in Guangdong province, South China. Mycology, 13(2), 143–152. https://doi.org/10.1080/21501203.2021.1978573
Mendu, L., Cobos, C. J., Tengey, T. K., Commey, L., Balasubramanian, V. K., Williams, L. D., Dhillon, K. K., Sharma, D., Pandey, M. K., Falalou, H., Varshney, R. K., Burow, M. D., Sudini, H. K., & Mendu, V. (2022). Seed coat mediated resistance against Aspergillus flavus infection in peanut. In Plant Gene (Vol. 32). Elsevier B.V. https://doi.org/10.1016/j.plgene.2022.100381
Mulyani, A., & Suwanda, M. H. (2019). Pengelolaan Lahan Kering Beriklim Kering untuk Pengembangan Jagung di Nusa Tenggara. https://doi.org/10.21082/jsdl.v13n1.2019.41-52
Page, M. J., McKenzie, J. E., Bossuyt, P. M., Boutron, I., Hoffmann, T. C., Mulrow, C. D., Shamseer, L., Tetzlaff, J. M., Akl, E. A., Brennan, S. E., Chou, R., Glanville, J., Grimshaw, J. M., Hróbjartsson, A., Lalu, M. M., Li, T., Loder, E. W., Mayo-Wilson, E., McDonald, S., … Moher, D. (2021). The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ, 372. https://doi.org/10.1136/BMJ.N71
Pandey, M. K., Kumar, R., Pandey, A. K., Soni, P., Gangurde, S. S., Sudini, H. K., Fountain, J. C., Liao, B., Desmae, H., Okori, P., Chen, X., Jiang, H., Mendu, V., Falalou, H., Njoroge, S., Mwololo, J., Guo, B., Zhuang, W., Wang, X., … Varshney, R. K. (2019). Mitigating aflatoxin contamination in groundnut through a combination of genetic resistance and post-harvest management practices. In Toxins (Vol. 11, Number 6). MDPI AG. https://doi.org/10.3390/toxins11060315
Pokhrel, S., Kharel, P., Pandey, S., Botton, S., Nugraha, G. T., Holbrook, C., & Ozias-Akins, P. (2024). Understanding the impacts of drought on peanuts (Arachis hypogaea L.): exploring physio-genetic mechanisms to develop drought-resilient peanut cultivars. In Frontiers in Genetics (Vol. 15). Frontiers Media SA. https://doi.org/10.3389/fgene.2024.1492434
Pollock, D., Davies, E. L., Peters, M. D. J., Tricco, A. C., Alexander, L., McInerney, P., Godfrey, C. M., Khalil, H., & Munn, Z. (2021). Undertaking a scoping review: A practical guide for nursing and midwifery students, clinicians, researchers, and academics. Journal of Advanced Nursing, 77(4), 2102–2113. https://doi.org/10.1111/jan.14743
Prasad, K., Yogendra, K., Sanivarapu, H., Rajasekaran, K., Cary, J. W., Sharma, K. K., & Bhatnagar-Mathur, P. (2023). Multiplexed Host-Induced Gene Silencing of Aspergillus flavus Genes Confers Aflatoxin Resistance in Groundnut. Toxins, 15(5). https://doi.org/10.3390/toxins15050319
Pushkarna, S., Gaba, K., Kharod, S., Kumar, A., Devi, S. U., Suneja, P., & Dang, A. S. (2025). Afatoxigenic Fungi: Human Health Impacts and Control Methods. Aflatoxigenic Fungi: Its Impact on Plant, Animal and Human Health, 197–208. https://doi.org/10.1201/9781003487043-25/aflatoxigenic-fungi-samridhi-pushkarna-kajal-gaba-shruti-kharod-anil-kumar-uma-devi-pooja-suneja-amita-suneja-dang
Rasheed, U., Cotty, P. J., Ain, Q. U., Wang, Y. F., & Liu, B. (2024). Efficacy of atoxigenic Aspergillus flavus from southern China as biocontrol agents against aflatoxin contamination in corn and peanuts. Pesticide Biochemistry and Physiology, 201. https://doi.org/10.1016/j.pestbp.2024.105887
Romero-Olivares, A. L., Lopez, A., Catalan-Dibene, J., Ferrenberg, S., Jordan, S. E., & Osborne, B. (2024). Effects of global change drivers on the expression of pathogenicity and stress genes in dryland soil fungi. MSphere, 9(11). https://doi.org/10.1128/msphere.00658-24
Safitri, A., Fauzi, M. T., & Hemon, A. F. (2026). Infeksi Aspergillus Flavus Pada Kacang Tanah Yang Diberi Gipsum Pada Kondisi Cekaman Kekeringan. Pascasarjana Universitas Mataram.
Safitri, A., Sarjan, M., Fauzi, T., & Permana, D. (2026). Journal of Multidisciplinary Science and Natural Resource Management Aplikasi Bioteknologi dalam Pengendalian Aspergillus flavus dan Mitigasi Kontaminasi Aflatoksin pada Kacang Tanah (Arachis hypogaea L.) Application of Biotechnology in Controlling Aspergillus flavus and Mitigating Aflatoxin Contamination in Peanuts (Arachis hypogaea L.). In Journal of Multidisciplinary Science and Natural Resource Management.
Singh, P., Callicott, K. A., Orbach, M. J., & Cotty, P. J. (2020). Molecular Analysis of S-morphology Aflatoxin Producers From the United States Reveals Previously Unknown Diversity and Two New Taxa. Frontiers in Microbiology, 11. https://doi.org/10.3389/fmicb.2020.01236
Sukhera, J. (2022). Narrative Reviews in Medical Education: Key Steps for Researchers. Journal of Graduate Medical Education, 14(4), 418–419. https://doi.org/10.4300/JGME-D-22-00481.1
Tumukunde, E., Xie, R., & Wang, S. (2021). Updates on the functions and molecular mechanisms of the genes involved in aspergillus flavus development and biosynthesis of aflatoxins. In Journal of Fungi (Vol. 7, Number 8). MDPI AG. https://doi.org/10.3390/jof7080666
Wang, P., Xu, J., Chang, P.-K., Liu, Z., & Kong, Q. (2022). New Insights of Transcriptional Regulator AflR in Aspergillus flavus Physiology. https://journals.asm.org/journal/spectrum
Wang, X., You, S. H., Lien, K. W., & Ling, M. P. (2019). Using disease-burden method to evaluate the strategies for reduction of aflatoxin exposure in peanuts. Toxicology Letters, 314, 75–81. https://doi.org/10.1016/J.TOXLET.2019.07.006
Wang, Y., Liu, D., Yin, H., Wang, H., Cao, C., Wang, J., Zheng, J., & Liu, J. (2023). Transcriptomic and Metabolomic Analyses of the Response of Resistant Peanut Seeds to Aspergillus flavus Infection. Toxins, 15(7). https://doi.org/10.3390/toxins15070414
Yu, B., Liu, N., Huang, L., Luo, H., Zhou, X., Lei, Y., Yan, L., Wang, X., Chen, W., Kang, Y., Ding, Y., Jin, G., Pandey, M. K., Janila, P., Kishan Sudini, H., Varshney, R. K., Jiang, H., Liu, S., & Liao, B. (2024). Identification and application of a candidate gene AhAftr1 for aflatoxin production resistance in peanut seed (Arachis hypogaea L.). Journal of Advanced Research, 62, 15–26. https://doi.org/10.1016/j.jare.2023.09.014
Zhao, X., Xing, M., Zhang, L. J., Han, S., Zhang, X., & Huang, J. (2025). Integrated metabolomic and transcriptomic analysis highlight the flavonoid response to antioxidant activity and Aspergillus flavus resistance in peanut seed coats. BMC Plant Biology, 25(1). https://doi.org/10.1186/s12870-025-07159-5
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