{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T05:19:47Z","timestamp":1787721587823,"version":"build-2784847793"},"reference-count":83,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T00:00:00Z","timestamp":1787529600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Henan Provincial University Science and Technology Innovation Team Support Program","award":["26IRTSTHN012"],"award-info":[{"award-number":["26IRTSTHN012"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32572360"],"award-info":[{"award-number":["32572360"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["32572360"],"award-info":[{"award-number":["32572360"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Outstanding Youth Science Fund of Henan Province","award":["242300421031"],"award-info":[{"award-number":["242300421031"]}]},{"name":"Natural Science Foundation of Henan province","award":["242300420471"],"award-info":[{"award-number":["242300420471"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Genes"],"abstract":"<jats:p>Background: Autophagy plays a central role in maintaining cellular homeostasis, regulating growth and development, and responding to multiple stresses. Autophagy-related genes (ATGs) play critical roles in autophagy, yet their functional diversity in soybean (Glycine max) remains underexplored. Methods: Genome-wide identification of GmATG genes was performed using sequence similarity and domain-based searches against the Wm82.gnm4 reference genome, followed by characterization of physicochemical properties, chromosomal distribution, phylogenetic relationships, gene duplication, conserved motifs, gene structure, three-dimensional structural, and promoter cis-acting elements. Tissue-specific expression and multiple stresses response were examined using transcriptome data and profiled by RT-qPCR. Results: A total of 60 GmATG genes belonging to 20 subfamilies were identified in soybean. Gene family expansion was predominantly driven by fragment duplication (33 gene pairs), with the ATG8 family expanding to 12 members, and pan-genomic analysis uncovered prominent copy number variation (6\u20139 copies) in the ATG18 family. GmATG genes showed distinct expression patterns in response to multiple abiotic and biotic stresses. Specifically, GmATG18f was significantly induced by phosphorus deficiency in the low-phosphorus-tolerant soybean variety Nannong 94-156. GmATG8g, GmATG9d and GmATG13d showed a typical expression trend of initial increase followed by decrease, with expression levels peaking at 6\u201312 h after salt stress treatment. GmATG8g and GmATG9d were rapidly upregulated at the early drought stress stage, while GmATG13a maintained sustained upregulation. In response to Phomopsis stem rot, GmATG7a\/8h\/8i\/11\/13d\/18e\/18f displayed differential expression in resistant and susceptible soybean materials. Conclusions: This study systematically characterizes the composition, expansion and stress response patterns of the GmATG gene family, revealing functional differentiation among family members. The identified key candidate genes, including abiotic-stress-regulated GmATG8g\/9d\/13d\/18f and biotic-stress-regulated GmATG7a\/8h\/8i\/11\/13d\/18e\/18f, provide valuable genetic resources for the molecular breeding of stress-tolerant soybean.<\/jats:p>","DOI":"10.3390\/genes17090996","type":"journal-article","created":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T17:20:28Z","timestamp":1787592028000},"page":"996","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Genome-Wide Identification of the GmATG Gene Family and Its Response to Multiple Biotic and Abiotic Stresses in Soybean (Glycine max)"],"prefix":"10.3390","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0009-0009-5488-8758","authenticated-orcid":false,"given":"Ling","family":"Yang","sequence":"first","affiliation":[{"name":"Collaborative Innovation Center of Henan Grain Crops, College of Agronomy, Henan Agricultural University, Zhengzhou 450002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-3236-1820","authenticated-orcid":false,"given":"Jingyi","family":"Fan","sequence":"additional","affiliation":[{"name":"Collaborative Innovation Center of Henan Grain Crops, College of Agronomy, Henan Agricultural University, Zhengzhou 450002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Enguang","family":"Ren","sequence":"additional","affiliation":[{"name":"Collaborative Innovation Center of Henan Grain Crops, College of Agronomy, Henan Agricultural University, Zhengzhou 450002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuo","family":"Yang","sequence":"additional","affiliation":[{"name":"Collaborative Innovation Center of Henan Grain Crops, College of Agronomy, Henan Agricultural University, Zhengzhou 450002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7204-9602","authenticated-orcid":false,"given":"Dandan","family":"Hu","sequence":"additional","affiliation":[{"name":"Collaborative Innovation Center of Henan Grain Crops, College of Agronomy, Henan Agricultural University, Zhengzhou 450002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,24]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1146\/annurev-arplant-042811-105441","article-title":"Autophagy: Pathways for self-eating in plant cells","volume":"63","author":"Liu","year":"2012","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1146\/annurev-arplant-042817-040606","article-title":"Autophagy: The Master of Bulk and Selective Recycling","volume":"69","author":"Marshall","year":"2018","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"e112534","DOI":"10.15252\/embj.2022112534","article-title":"Selective autophagy regulates chloroplast protein import and promotes plant stress tolerance","volume":"42","author":"Wan","year":"2023","journal-title":"EMBO J."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1016\/S1534-5807(03)00296-X","article-title":"A unified nomenclature for yeast autophagy-related genes","volume":"5","author":"Klionsky","year":"2003","journal-title":"Dev. Cell"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1146\/annurev-cellbio-092910-154005","article-title":"The role of Atg proteins in autophagosome formation","volume":"27","author":"Mizushima","year":"2011","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"33105","DOI":"10.1074\/jbc.M204630200","article-title":"The APG8\/12-activating enzyme APG7 is required for proper nutrient recycling and senescence in Arabidopsis thaliana","volume":"277","author":"Doelling","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.pbi.2005.01.013","article-title":"Autophagic recycling: Lessons from yeast help define the process in plants","volume":"8","author":"Thompson","year":"2005","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1007\/s13238-011-1104-4","article-title":"Role of plant autophagy in stress response","volume":"2","author":"Han","year":"2011","journal-title":"Protein Cell"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1093\/dnares\/dsr024","article-title":"Genome-wide identification, classification, and expression analysis of autophagy-associated gene homologues in rice (Oryza sativa L.)","volume":"18","author":"Xia","year":"2011","journal-title":"DNA Res."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"263803","DOI":"10.1242\/jcs.263803","article-title":"An A. thaliana mutant lacking all nine ATG8 isoforms provides genetic evidence for functional specialization of ATG8 in plants","volume":"138","author":"Grujic","year":"2025","journal-title":"J. Cell Sci."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Wang, Z., Luo, F., Meng, C., Si, J., Zhang, W., Sun, J., and Wang, Z. (2025). Identification of the ZmATG18 subfamily genes in maize and the role of ZmATG18a in drought stress. BMC Genom., 26.","DOI":"10.1186\/s12864-025-11910-5"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1105\/tpc.19.00413","article-title":"Arabidopsis SINAT Proteins Control Autophagy by Mediating Ubiquitylation and Degradation of ATG13","volume":"32","author":"Qi","year":"2020","journal-title":"Plant Cell"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Luo, L., Zhang, P., Zhu, R., Fu, J., Su, J., Zheng, J., Wang, Z., Wang, D., and Gong, Q. (2017). Autophagy Is Rapidly Induced by Salt Stress and Is Required for Salt Tolerance in Arabidopsis. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.01459"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"3915","DOI":"10.1093\/jxb\/eru008","article-title":"Delayed degradation of chlorophylls and photosynthetic proteins in Arabidopsis autophagy mutants during stress-induced leaf yellowing","volume":"65","author":"Sakuraba","year":"2014","journal-title":"J. Exp. Bot."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"101200","DOI":"10.1016\/j.stress.2025.101200","article-title":"Differential induction of autophagy and miRNA-mediated regulation modulate salt tolerance in contrasting rice varieties","volume":"19","author":"Talakayala","year":"2026","journal-title":"Plant Stress"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1681","DOI":"10.1038\/s41477-025-02057-y","article-title":"Plasma membrane CYBDOM proteins catalyse apoplastic AsA regeneration and interact with RbohD to activate autophagy and drought tolerance in plants","volume":"11","author":"Chen","year":"2025","journal-title":"Nat. Plants"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Cheng, S., Fan, S., Yang, C., Hu, W., and Liu, F. (2025). Proteomics revealed novel functions and drought tolerance of Arabidopsis thaliana protein kinase ATG1. BMC Biol., 23.","DOI":"10.1186\/s12915-025-02149-3"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"5889","DOI":"10.1038\/s41467-025-61191-5","article-title":"The N-degron pathway governs autophagy to promote thermotolerance in Arabidopsis","volume":"16","author":"Kim","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"10910","DOI":"10.1038\/s41467-024-55292-w","article-title":"Stress granules sequester autophagy proteins to facilitate plant recovery from heat stress","volume":"15","author":"Li","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Yu, J., Zhen, X., Li, X., Li, N., and Xu, F. (2019). Increased Autophagy of Rice Can Increase Yield and Nitrogen Use Efficiency (NUE). Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.00584"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Zhen, X., Xu, F., Zhang, W., Li, N., and Li, X. (2019). Overexpression of rice gene OsATG8b confers tolerance to nitrogen starvation and increases yield and nitrogen use efficiency (NUE) in Arabidopsis. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0223011"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"uhac068","DOI":"10.1093\/hr\/uhac068","article-title":"Autophagic pathway contributes to low-nitrogen tolerance by optimizing nitrogen uptake and utilization in tomato","volume":"9","author":"Cao","year":"2022","journal-title":"Hortic. Res."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"460","DOI":"10.1104\/pp.18.01379","article-title":"The Local Phosphate Deficiency Response Activates Endoplasmic Reticulum Stress-Dependent Autophagy","volume":"179","author":"Naumann","year":"2019","journal-title":"Plant Physiol."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Yoshitake, Y., and Yoshimoto, K. (2022). Intracellular phosphate recycling systems for survival during phosphate starvation in plants. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.1088211"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"102781","DOI":"10.1016\/j.pbi.2025.102781","article-title":"The multifaceted roles of autophagy in plant immunity","volume":"87","author":"Jeon","year":"2025","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1016\/j.cell.2009.02.036","article-title":"Autophagic components contribute to hypersensitive cell death in Arabidopsis","volume":"137","author":"Hofius","year":"2009","journal-title":"Cell"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"818","DOI":"10.1111\/j.1365-313X.2011.04546.x","article-title":"Autophagy differentially controls plant basal immunity to biotrophic and necrotrophic pathogens","volume":"66","author":"Lenz","year":"2011","journal-title":"Plant J. Cell Mol. Biol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"112577","DOI":"10.1016\/j.plantsci.2025.112577","article-title":"Research progress on the autophagy gene ATG6 in planta","volume":"359","author":"Wang","year":"2025","journal-title":"Plant Sci."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Tang, C.X., Liu, T.T., Ge, Q.D., Liu, X.Y., Wang, W., Sun, Y., Liu, Y., Li, F., and Wang, G.F. (2026). The BTB\/POZ-MATH Protein ZmBPM1 Interacts With Autophagy-related Protein ZmATG6 and Modulates the NLR Protein Rp1-D21-Mediated Defense Response in Maize. Adv. Sci., e77193.","DOI":"10.1002\/advs.77193"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"eadu9554","DOI":"10.1126\/science.adu9554","article-title":"Selective autophagy fine-tunes plant immunity to promote cell survival during viral infection","volume":"392","author":"Clavel","year":"2026","journal-title":"Science"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"10918","DOI":"10.1021\/acs.jafc.6c00361","article-title":"Plant Virus and Vector Insect Regulate the Dual Phosphorylation of ATG1\u2013ATG13 To Maintain a Moderate Autophagy for Viral Persistent Propagation","volume":"74","author":"Liu","year":"2026","journal-title":"J. Agric. Food Chem."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Xia, T., Xiao, D., Liu, D., Chai, W., Gong, Q., and Wang, N.N. (2012). Heterologous expression of ATG8c from soybean confers tolerance to nitrogen deficiency and increases yield in Arabidopsis. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0037217"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/j.jplph.2016.01.007","article-title":"Nitrogen redistribution and its relationship with the expression of GmATG8c during seed filling in soybean","volume":"192","author":"Islam","year":"2016","journal-title":"J. Plant Physiol."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Hashimi, S.M., Wu, N.N., Ran, J., and Liu, J.Z. (2021). Silencing Autophagy-Related Gene 2 (ATG2) Results in Accelerated Senescence and Enhanced Immunity in Soybean. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms222111749"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Hashimi, S.M., Huang, M.J., Amini, M.Q., Wang, W.X., Liu, T.Y., Chen, Y., Liao, L.N., Lan, H.J., and Liu, J.Z. (2023). Silencing GmATG7 Leads to Accelerated Senescence and Enhanced Disease Resistance in Soybean. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms242216508"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1038\/nature08670","article-title":"Genome sequence of the palaeopolyploid soybean","volume":"463","author":"Schmutz","year":"2010","journal-title":"Nature"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"D1496","DOI":"10.1093\/nar\/gkaa1107","article-title":"A new decade and new data at SoyBase, the USDA-ARS soybean genetics and genomics database","volume":"49","author":"Brown","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1194","DOI":"10.1016\/j.molp.2020.06.009","article-title":"TBtools: An Integrative Toolkit Developed for Interactive Analyses of Big Biological Data","volume":"13","author":"Chen","year":"2020","journal-title":"Mol. Plant"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1093\/bioinformatics\/bti125","article-title":"Protein homology detection by HMM-HMM comparison","volume":"21","year":"2005","journal-title":"Bioinformatics"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"D733","DOI":"10.1093\/nar\/gkv1189","article-title":"Reference sequence (RefSeq) database at NCBI: Current status, taxonomic expansion, and functional annotation","volume":"44","author":"Wright","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"W585","DOI":"10.1093\/nar\/gkm259","article-title":"WoLF PSORT: Protein localization predictor","volume":"35","author":"Horton","year":"2007","journal-title":"Nucleic Acids Res."},{"key":"ref_42","first-page":"671","article-title":"Application analysis of predictors for plant protein subcellular localization based on proteome data of Camellia sinensis (L.) O. Ktze","volume":"38","author":"Liu","year":"2020","journal-title":"Plant Sci. J."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1093\/molbev\/msu300","article-title":"IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies","volume":"32","author":"Nguyen","year":"2015","journal-title":"Mol. Biol. Evol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"D1202","DOI":"10.1093\/nar\/gkr1090","article-title":"The Arabidopsis Information Resource (TAIR): Improved gene annotation and new tools","volume":"40","author":"Lamesch","year":"2012","journal-title":"Nucleic Acids Res."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"e3","DOI":"10.1093\/pcp\/pcu183","article-title":"Triticeae resources in Ensembl Plants","volume":"56","author":"Bolser","year":"2015","journal-title":"Plant Cell Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"794","DOI":"10.1016\/j.molp.2023.03.011","article-title":"SoyOmics: A deeply integrated database on soybean multi-omics","volume":"16","author":"Liu","year":"2023","journal-title":"Mol. Plant"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1016\/j.cell.2020.05.023","article-title":"Pan-Genome of Wild and Cultivated Soybeans","volume":"182","author":"Liu","year":"2020","journal-title":"Cell"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Nystrom, S.L., and McKay, D.J. (2021). Memes: A motif analysis environment in R using tools from the MEME Suite. PLoS Comput. Biol., 17.","DOI":"10.1371\/journal.pcbi.1008991"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"W296","DOI":"10.1093\/nar\/gky427","article-title":"SWISS-MODEL: Homology modelling of protein structures and complexes","volume":"46","author":"Waterhouse","year":"2018","journal-title":"Nucleic Acids Res."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1093\/nar\/30.1.325","article-title":"PlantCARE, a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences","volume":"30","author":"Lescot","year":"2002","journal-title":"Nucleic Acids Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"669","DOI":"10.1016\/j.molp.2025.02.003","article-title":"A large-scale integrated transcriptomic atlas for soybean organ development","volume":"18","author":"Fan","year":"2025","journal-title":"Mol. Plant"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1016\/j.cell.2024.10.050","article-title":"A spatially resolved multi-omic single-cell atlas of soybean development","volume":"188","author":"Zhang","year":"2025","journal-title":"Cell"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2176","DOI":"10.1093\/plcell\/koae041","article-title":"The Myb73-GDPD2-GA2ox1 transcriptional regulatory module confers phosphate deficiency tolerance in soybean","volume":"36","author":"Hu","year":"2024","journal-title":"Plant Cell"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2362","DOI":"10.1111\/pbi.13668","article-title":"Nuclear factor Y subunit GmNFYA competes with GmHDA13 for interaction with GmFVE to positively regulate salt tolerance in soybean","volume":"19","author":"Lu","year":"2021","journal-title":"Plant Biotechnol. J."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Le, D.T., Nishiyama, R., Watanabe, Y., Tanaka, M., Seki, M., Ham Le, H., Yamaguchi-Shinozaki, K., Shinozaki, K., and Tran, L.S. (2012). Differential gene expression in soybean leaf tissues at late developmental stages under drought stress revealed by genome-wide transcriptome analysis. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0049522"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"112622","DOI":"10.1016\/j.plantsci.2025.112622","article-title":"Functional analysis of GmMATE gene family in soybean phosphorus homeostasis and abiotic stress resilience","volume":"359","author":"Xu","year":"2025","journal-title":"Plant Sci."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) Method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1578","DOI":"10.1080\/15548627.2025.2469189","article-title":"Establishment of a yeast essential protein conditional-degradation library and screening for autophagy-regulating genes","volume":"21","author":"Zhang","year":"2025","journal-title":"Autophagy"},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Chen, H., Dong, J., and Wang, T. (2021). Autophagy in Plant Abiotic Stress Management. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22084075"},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Huang, S., Zhang, B., and Chen, W. (2021). Research Progress of ATGs Involved in Plant Immunity and NPR1 Metabolism. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms222212093"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"22933","DOI":"10.1038\/s41598-021-02239-6","article-title":"Genome-wide analysis of autophagy-related genes in Medicago truncatula highlights their roles in seed development and response to drought stress","volume":"11","author":"Yang","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Quezada-Rodriguez, E.H., Gomez-Velasco, H., Arthikala, M.K., Lara, M., Hernandez-Lopez, A., and Nanjareddy, K. (2021). Exploration of Autophagy Families in Legumes and Dissection of the ATG18 Family with a Special Focus on Phaseolus vulgaris. Plants, 10.","DOI":"10.3390\/plants10122619"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1093\/pcp\/pcad015","article-title":"Autophagy-Mediated Phosphate Homeostasis in Arabidopsis Involves Modulation of Phosphate Transporters","volume":"64","author":"Chiu","year":"2023","journal-title":"Plant Cell Physiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"886","DOI":"10.1093\/plphys\/kiad133","article-title":"Autophagy and multivesicular body pathways cooperate to protect sulfur assimilation and chloroplast functions","volume":"192","author":"Fu","year":"2023","journal-title":"Plant Physiol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"113398","DOI":"10.1016\/j.ecoenv.2022.113398","article-title":"The protective role of MdATG10-mediated autophagy in apple plant under cadmium stress","volume":"234","author":"Huo","year":"2022","journal-title":"Ecotox. Environ. Saf."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"107752","DOI":"10.1016\/j.isci.2023.107752","article-title":"The PtdIns3P phosphatase MtMP promotes symbiotic nitrogen fixation via mitophagy in Medicago truncatula","volume":"26","author":"Xue","year":"2023","journal-title":"iScience"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"3936","DOI":"10.1093\/plcell\/koac185","article-title":"Defects in autophagy lead to selective in vivo changes in turnover of cytosolic and organelle proteins in Arabidopsis","volume":"34","author":"Li","year":"2022","journal-title":"Plant Cell"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"2466","DOI":"10.1111\/nph.70564","article-title":"P3IPs activate autophagy by disrupting the GAPC2\u2013ATG3 interaction and target TuMV 6K2 for degradation","volume":"248","author":"Chen","year":"2025","journal-title":"New Phytol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"101868","DOI":"10.1016\/j.xplc.2026.101868","article-title":"Antiviral RNAi defective 4 (AVI4) encodes a lysophospholipid acyltransferase coopting with AVI2 and ATG8a to promote antiviral immunity in Arabidopsis","volume":"7","author":"Yan","year":"2026","journal-title":"Plant Commun."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"100937","DOI":"10.1016\/j.xplc.2024.100937","article-title":"Clathrin light chains negatively regulate plant immunity by hijacking the autophagy pathway","volume":"5","author":"Lan","year":"2024","journal-title":"Plant Commun."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"2093","DOI":"10.1080\/15548627.2020.1810426","article-title":"Phosphorylation of ATG18a by BAK1 suppresses autophagy and attenuates plant resistance against necrotrophic pathogens","volume":"17","author":"Zhang","year":"2021","journal-title":"Autophagy"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1472","DOI":"10.1016\/j.molp.2025.07.006","article-title":"OsATG1 and OsATG8 exhibit autophagy-independent functions to oppositely regulate ROP GTPase-mediated plant immunity in rice","volume":"18","author":"He","year":"2025","journal-title":"Mol. Plant"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"2018","DOI":"10.1111\/pbi.70403","article-title":"TaBZR1 Directly Activates Autophagy-Related (ATG) 8 to Promote Wheat Defence to Powdery Mildew","volume":"24","author":"Zhuang","year":"2026","journal-title":"Plant Biotechnol. J."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"4769","DOI":"10.1038\/s41467-023-40472-x","article-title":"Autophagy promotes jasmonate-mediated defense against nematodes","volume":"14","author":"Zou","year":"2023","journal-title":"Nat. Commun."},{"key":"ref_75","first-page":"1","article-title":"Over expression of ATG8i enhances disease resistance in Arabidopsis","volume":"61","author":"Wang","year":"2026","journal-title":"Chin. Bull. Bot."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1745","DOI":"10.1080\/15548627.2022.2153573","article-title":"The Valsa Mali effector Vm1G-1794 protects the aggregated MdEF-Tu from autophagic degradation to promote infection in apple","volume":"19","author":"Che","year":"2023","journal-title":"Autophagy"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Zhang, Y.L., Yang, Z.K., Zhang, Z., Wang, G.P., Li, X.D., and Hong, N. (2025). Citrus tristeza virus p20 suppresses antiviral RNA silencing by co-opting autophagy-related protein 8 to mediate the autophagic degradation of SGS3. PLoS Pathog., 21.","DOI":"10.1371\/journal.ppat.1012960"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"2891","DOI":"10.1111\/pbi.70119","article-title":"A fijivirus capsid protein hijacks autophagy degrading an \u03c9-3 fatty acid desaturase to suppress jasmonate-mediated antiviral defence","volume":"23","author":"Liu","year":"2025","journal-title":"Plant Biotechnol. J."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.tplants.2025.07.010","article-title":"Phosphorus acquisition and pathogen defense: Synergies versus trade-offs","volume":"31","author":"Ding","year":"2026","journal-title":"Trends Plant Sci."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"1068","DOI":"10.1111\/jipb.13662","article-title":"Autophagy receptor ZmNBR1 promotes the autophagic degradation of ZmBRI1a and enhances drought tolerance in maize","volume":"66","author":"Xiang","year":"2024","journal-title":"J. Integr. Plant Biol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"2033","DOI":"10.1080\/15548627.2015.1098798","article-title":"Tomato HsfA1a plays a critical role in plant drought tolerance by activating ATG genes and inducing autophagy","volume":"11","author":"Wang","year":"2015","journal-title":"Autophagy"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"862","DOI":"10.1080\/15548627.2019.1643656","article-title":"Dehydrin MtCAS31 promotes autophagic degradation under drought stress","volume":"16","author":"Li","year":"2020","journal-title":"Autophagy"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"954","DOI":"10.4161\/auto.5.7.9290","article-title":"Autophagy is required for tolerance of drought and salt stress in plants","volume":"5","author":"Liu","year":"2009","journal-title":"Autophagy"}],"container-title":["Genes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4425\/17\/9\/996\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T04:26:58Z","timestamp":1787718418000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4425\/17\/9\/996"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,24]]},"references-count":83,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["genes17090996"],"URL":"https:\/\/doi.org\/10.3390\/genes17090996","relation":{},"ISSN":["2073-4425"],"issn-type":[{"value":"2073-4425","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,24]]}}}