{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T12:58:32Z","timestamp":1790600312645,"version":"4.1.0"},"reference-count":87,"publisher":"MDPI AG","issue":"19","license":[{"start":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T00:00:00Z","timestamp":1790553600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100012901","name":"Zhoukou Normal University","doi-asserted-by":"publisher","award":["ZKNUC2023081"],"award-info":[{"award-number":["ZKNUC2023081"]}],"id":[{"id":"10.13039\/100012901","id-type":"DOI","asserted-by":"publisher"}]},{"award":["ZKNUC2023081"],"award-info":[{"award-number":["ZKNUC2023081"]}],"id":[{"id":"https:\/\/ror.org\/00jjkh886","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Science and Technology Research Project of Henan Province","award":["262102110178"],"award-info":[{"award-number":["262102110178"]}]},{"name":"Central Guiding Local Science and Technology Development Fund Project in Henan Province","award":["Z20251831182"],"award-info":[{"award-number":["Z20251831182"]}]},{"name":"Key Research and Development Project of Henan Province","award":["251111110900"],"award-info":[{"award-number":["251111110900"]}]},{"name":"Henan Province Science and Technology Research and Development Joint Fund (Industry) Major Project","award":["235101610015"],"award-info":[{"award-number":["235101610015"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Wheat production is severely constrained by Fusarium crown rot (FCR) caused by Fusarium pseudograminearum and low soil nutrient use efficiency, making biocontrol an important strategy for sustainable agriculture. This study characterized a newly isolated Trichoderma harzianum strain, M2, and explored its bioprotective and growth-promoting mechanisms in wheat. Identified via morphological and phylogenetic analyses, strain M2 exhibited 91.24% in vitro inhibition against pathogenic F. pseudograminearum. Its 105 cfu\/mL spore suspension remodeled root architecture and increased seedling biomass in hydroponic and pot experiments. Greenhouse and field assays recorded FCR control efficacies of 90.12% and 71.70%, respectively; field application of M2 reduced the disease index from 82% to 28%. RNA-seq and qRT-PCR indicated that M2 confers FCR resistance by triggering induced systemic resistance. Microscopy demonstrated that M2 penetrates root epidermal cells, forms biofilm-like structures and stably colonizes root xylem tissues. Furthermore, M2 enhanced soil phosphorus uptake and phosphorus accumulation in wheat. Collectively, endophytic root colonization and multi-hormone regulatory pathways enable T. harzianum M2 to improve wheat defense and performance, representing a promising agent for eco-friendly wheat production.<\/jats:p>","DOI":"10.3390\/plants15192957","type":"journal-article","created":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T11:35:59Z","timestamp":1790595359000},"page":"2957","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Trichoderma harzianum M2 Induces Systemic Resistance to Fusarium Crown Rot and Remodels Root Architecture to Promote Phosphorus Uptake in Wheat"],"prefix":"10.3390","volume":"15","author":[{"given":"Junchang","family":"Li","sequence":"first","affiliation":[{"name":"Henan Province Key Laboratory of Efficient Crop Production and Food Quality Safety, Zhoukou Normal University, Zhoukou 466001, China"},{"name":"Modern Agricultural Industry Research Institute, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"conceptualization","vocabulary":"credit"}]},{"given":"Yingying","family":"Jin","sequence":"additional","affiliation":[{"name":"Modern Agricultural Industry Research Institute, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"},{"name":"Wheat Technology Innovation Center, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"}]},{"given":"Yingxue","family":"Wang","sequence":"additional","affiliation":[{"name":"Modern Agricultural Industry Research Institute, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"},{"name":"Wheat Technology Innovation Center, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"}]},{"given":"Yiying","family":"Chen","sequence":"additional","affiliation":[{"name":"Modern Agricultural Industry Research Institute, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"},{"name":"Wheat Technology Innovation Center, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"}]},{"given":"Tingyu","family":"Wang","sequence":"additional","affiliation":[{"name":"Modern Agricultural Industry Research Institute, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"},{"name":"Wheat Technology Innovation Center, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"}]},{"given":"Xingdong","family":"Yang","sequence":"additional","affiliation":[{"name":"Henan Province Key Laboratory of Efficient Crop Production and Food Quality Safety, Zhoukou Normal University, Zhoukou 466001, China"},{"name":"Modern Agricultural Industry Research Institute, Henan Zhoukou Agricultural High-Tech Industry Demonstration Zone, Zhoukou 477100, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"visualization","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1038\/nature11650","article-title":"Analysis of the bread wheat genome using whole-genome shotgun sequencing","volume":"491","author":"Brenchley","year":"2012","journal-title":"Nature"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"20260","DOI":"10.1073\/pnas.1116437108","article-title":"Global food demand and the sustainable intensification of agriculture","volume":"108","author":"Tilman","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"5411","DOI":"10.1093\/jxb\/ert333","article-title":"Hybrid breeding in wheat: Technologies to improve hybrid wheat seed production","volume":"64","author":"Whitford","year":"2013","journal-title":"J. Exp. Bot."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"756","DOI":"10.1016\/j.foodcont.2013.06.019","article-title":"Mycotoxin glucosylation in commercial wheat varieties: Impact on resistance to Fusarium graminearum under laboratory and field conditions","volume":"34","author":"Audenaert","year":"2013","journal-title":"Food Control"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1547","DOI":"10.1111\/mpp.12639","article-title":"Fusarium crown rot caused by Fusarium pseudograminearum in cereal crops: Recent progress and future prospects","volume":"19","author":"Kazan","year":"2018","journal-title":"Mol. Plant Pathol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"769","DOI":"10.1023\/A:1026042711064","article-title":"Association of Fusarium species in the wheat stem rot complex","volume":"109","author":"Pettitt","year":"2003","journal-title":"Eur. J. Plant Pathol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"416","DOI":"10.1080\/07060660209507029","article-title":"Insights into the prevalence and management of soilborne cereal pathogens under direct seeding in the Pacific Northwest, USA","volume":"24","author":"Paulitz","year":"2002","journal-title":"Can. J. Plant Pathol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.1111\/ppa.12821","article-title":"Development of three Fusarium crown rot causal agents and systemic translocation of deoxynivalenol following stem base infection of soft wheat","volume":"67","author":"Beccari","year":"2018","journal-title":"Plant Pathol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1111\/pbr.12274","article-title":"Resistance to Fusarium crown rot in wheat and barley: A review","volume":"134","author":"Liu","year":"2015","journal-title":"Plant Breed."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1038\/s41579-022-00819-5","article-title":"Trichoderma: A multipurpose, plant-beneficial microorganism for eco-sustainable agriculture","volume":"21","author":"Woo","year":"2023","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Hinterdobler, W., Li, G., Spiegel, K., Basyouni-Khamis, S., Gorfer, M., and Schmoll, M. (2021). Trichoderma reesei isolated from Austrian soil with high potential for biotechnological application. Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.552301"},{"key":"ref_12","first-page":"966","article-title":"Application andmechanism of Trichoderma in biological control of plant disease","volume":"35","author":"You","year":"2019","journal-title":"Chin. J. Biol. Control"},{"key":"ref_13","first-page":"279","article-title":"Induced systemic resistance in Trichoderma harzianum T39 biocontrol of Botrytis cinerea","volume":"104","author":"Bigirimana","year":"1988","journal-title":"Eur. J. Plant Pathol."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Abdelkhalek, A., Al-Askar, A., Arishi, A., and Behiry, S. (2022). Trichoderma hamatum strain Th23 promotes tomato growth and induces systemic resistance against tobacco mosaic virus. J. Fungi, 8.","DOI":"10.3390\/jof8030228"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1038\/nrmicro797","article-title":"Trichoderma species-opportunistic, avirulent plant symbionts","volume":"2","author":"Harman","year":"2004","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/j.biocontrol.2015.10.006","article-title":"Integrated management of Fusarium crown rot of wheat usingfungicide seed treatment, cultivar resistance, and induction of systemic acquired resistance (SAR)","volume":"92","author":"Moyaelizondo","year":"2016","journal-title":"Biol. Control"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"101475","DOI":"10.1016\/j.pmpp.2020.101475","article-title":"The application potential of Trichoderma T-soybean containing 1-aminocyclopropane-1-carboxylate for maize production","volume":"110","author":"Zhang","year":"2020","journal-title":"Physiol. Mol. Plant Pathol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"126767","DOI":"10.1016\/j.micres.2021.126767","article-title":"Trichoderma asperellum xylanases promote growth and induce resistance in poplar","volume":"248","author":"Guo","year":"2021","journal-title":"Microbiol. Res."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Yao, X., Guo, H., Zhang, K., Zhao, M., Ruan, J., and Chen, J. (2023). Trichoderma and its role in biological control of plant fungal and nematode disease. Front. Microbiol., 14.","DOI":"10.3389\/fmicb.2023.1160551"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Vitti, A., Pellegrini, E., Nali, C., Lovelli, S., Sofo, A., Valerio, M., Scopa, A., and Nuzzaci, M. (2016). Trichoderma harzianum T-22 induces systemic resistance in tomato infected by Cucumber mosaic virus. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01520"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1186\/s41938-023-00728-6","article-title":"Potential of endophytic fungi as a pathogenic biocontrol agent and growth promoters in corn seedlings","volume":"33","author":"Fauriah","year":"2023","journal-title":"Egypt. J. Biol. Pest Control"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"You, J., Li, G., Li, C., Zhu, L., Yang, H., Song, R., and Gu, W. (2022). Biological control and plant growth promotion by volatile organic compounds of Trichoderma koningiopsis T-51. J. Fungi, 8.","DOI":"10.3390\/jof8020131"},{"key":"ref_23","first-page":"13","article-title":"The effect of Trichoderma harziaiarum H-13 on the rice nitrate reductase activity and N, P, K content","volume":"30","author":"Huang","year":"2003","journal-title":"Microbiol. China"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"e1176654","DOI":"10.1080\/15592324.2016.1176654","article-title":"Development of marker genes for jasmonic acid signaling in shoots and roots of wheat","volume":"11","author":"Liu","year":"2016","journal-title":"Plant Signal. Behav."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1146\/annurev-phyto-073009-114447","article-title":"Hormone crosstalk in plant disease and defense: More than just jasmonate-salicylate antagonism","volume":"49","author":"Grant","year":"2011","journal-title":"Annu. Rev. Phytopathol."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Veselova, S., Nuzhnaya, T., Burkhanova, G., Rumyantsev, S., Khusnutdinova, E., and Maksimov, I. (2021). Ethylene-cytokinin interaction determines early defense response of wheat against Stagonospora nodorum berk. Biomolecules, 11.","DOI":"10.3390\/biom11020174"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"6266","DOI":"10.1038\/s41467-020-20133-z","article-title":"Cloning of wheat keto-acyl thiolase 2B reveals a role of jasmonic acid in grain weight determination","volume":"11","author":"Chen","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1007\/s00122-023-04289-y","article-title":"Genome-wide associa tion analysis of Fusarium crown rot resistance in Chinese wheat landraces","volume":"136","author":"Hou","year":"2023","journal-title":"Theor. Appl. Genet."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Xu, F., Yang, G., Wang, J., Song, Y., Liu, L., Zhao, K., Li, Y., and Han, Z. (2018). Spatial distribution of root and crown rot fungi associated with winter wheat in the North China Plain and its relationship with climate variables. Front. Microbiol., 9.","DOI":"10.3389\/fmicb.2018.01054"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1016\/S1055-7903(02)00400-1","article-title":"Multilocus phylogenetic structure within the Trichoderma harzianum\/Hypocrea lixii complex","volume":"27","author":"Chaverri","year":"2003","journal-title":"Mol. Phylogenet. Evol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1080\/15572536.2003.11833257","article-title":"Trichoderma species associated with the green mold epidemic of commercially grown Agaricus bisporus","volume":"94","author":"Samuels","year":"2002","journal-title":"Mycologia"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1094\/PBIOMES-08-22-0056-R","article-title":"Biocontrol impacts on wheat physiology and Fusarium head blight outcomes are bacterial endophyte strain and cultivar specific","volume":"7","author":"Whitaker","year":"2023","journal-title":"Phytobiomes J."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"126589","DOI":"10.1016\/j.micres.2020.126589","article-title":"Interkingdom signaling in plant-rhizomicrobiome interactions for sustainable agriculture","volume":"241","author":"Phour","year":"2020","journal-title":"Microbiol. Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.soilbio.2007.07.002","article-title":"Trichoderma-plant-pathogen interactions","volume":"40","author":"Vinale","year":"2008","journal-title":"Soil Biol. Biochem."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"128719","DOI":"10.1016\/j.micres.2026.128719","article-title":"Trichoderma specialized metabolites in biocontrol: Gene\u2013metabolite links, ecological functions, and translational bottlenecks","volume":"314","author":"Riolo","year":"2026","journal-title":"Microbiol. Res."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1016\/j.fbr.2021.12.004","article-title":"Role and genetic basis of specialised secondary metabolites in Trichoderma ecophysiology","volume":"39","author":"Vicente","year":"2022","journal-title":"Fungal Biol. Rev."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1367","DOI":"10.1039\/D5NP00017C","article-title":"Advances in the discovery and study of Trichoderma natural products for biological control applications","volume":"42","author":"Jin","year":"2025","journal-title":"Nat. Prod. Rep."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1007\/s42360-026-00969-z","article-title":"Trichoderma species as sustainable biocontrol agents: Mechanisms, applications, and future prospects in plant disease management","volume":"79","author":"Kumar","year":"2026","journal-title":"Indian Phytopathol."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Tiwari, M., Singh, R., Jha, R., and Singh, P. (2022). Heritable priming by Trichoderma: A sustainable approach for wheat protection against Bipolaris sorokiniana. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.1050765"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"654","DOI":"10.1111\/jam.12387","article-title":"Trichoderma harzianum elicits induced resistance in sunflower challenged by Rhizoctonia solani","volume":"116","author":"Singh","year":"2014","journal-title":"J. Appl. Microbiol."},{"key":"ref_41","first-page":"4162","article-title":"Preliminary study on the control effect of Trichoderma harzianum EMF910 on the root rot pathogens of Astragalus membranaceus in Ningxia saline-alkali regions","volume":"51","author":"Zhang","year":"2024","journal-title":"Microbiol. China"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Niu, J., Yan, X., Bai, Y., Li, W., Lu, G., Wang, Y., Liu, H., Shi, Z., and Liang, J. (2024). Integration of transcriptomics and WGCNA to characterize Trichoderma harzianum-induced systemic resistance in Astragalus mongholicus for defense against Fusarium solani. Genes, 15.","DOI":"10.3390\/genes15091180"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"127182","DOI":"10.1016\/j.micres.2022.127182","article-title":"Biocontrol potential of Trichoderma harzianum CGMCC20739 (Tha739) against postharvest bitter rot of apples","volume":"265","author":"Zhang","year":"2022","journal-title":"Microbiol. Res."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Wang, G., and Zeng, H. (2022). Antibacterial effect of cell-free supernatant from Lactobacillus pentosus L-36 against Staphylococcus aureus from bovine mastitis. Molecules, 27.","DOI":"10.3390\/molecules27217627"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1080\/03235408.2016.1263052","article-title":"Alleviation of Fusarium oxysporum induced oxidative stress in wheat by Trichoderma viride","volume":"50","author":"Mohapatra","year":"2017","journal-title":"Arch. Phytopath. Plant"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Zhang, Y., Lu, Y., Jin, Z., Li, B., Wu, L., and He, Y. (2024). Antifungal mechanism of cell-free supernatant produced by Trichoderma virens and its efficacy for the control of pear Valsa canker. Front. Microbiol., 15.","DOI":"10.3389\/fmicb.2024.1377683"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1899","DOI":"10.1111\/nph.19935","article-title":"A potent endophytic fungus Purpureocillium lilacinum YZ1 protects against Fusarium infection in field-grown wheat","volume":"243","author":"Kimotho","year":"2024","journal-title":"New Phytol."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Guti\u00e9rrez-Moreno, K., Olgu\u00edn-Mart\u00ednez, A., Montoya-Mart\u00ednez, A., and de los Santos-Villalobos, S. (2025). Trichoderma in sustainable agriculture and the challenges related to its effectiveness. Diversity, 17.","DOI":"10.3390\/d17100734"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"e70205","DOI":"10.1111\/jph.70205","article-title":"Underground warfare: Mechanisms of Trichoderma in the suppression of the march of Fusarium in banana plants\u2014Advances, limitations and perspectives","volume":"173","year":"2025","journal-title":"J. Phytopathol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1093\/femsre\/fuad066","article-title":"Root colonization by beneficial rhizobacteria","volume":"48","author":"Liu","year":"2024","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1146\/annurev.micro.62.081307.162918","article-title":"Plant-growth-promoting rhizobacteria","volume":"63","author":"Lugtenberg","year":"2009","journal-title":"Annu. Rev. Microbiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1146\/annurev-phyto-073009-114450","article-title":"Induced systemic resistance and plant responses to fungal biocontrol agents","volume":"48","author":"Shoresh","year":"2010","journal-title":"Annu. Rev. Phytopathol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"749","DOI":"10.1038\/nrmicro2637","article-title":"Trichoderma: The genomics of opportunistic success","volume":"9","author":"Druzhinina","year":"2011","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_54","first-page":"524","article-title":"Priming: Getting ready for battle","volume":"27","author":"Conrath","year":"2006","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1038\/nature22009","article-title":"Plant signalling in symbiosis and immunity","volume":"543","author":"Zipfel","year":"2017","journal-title":"Nature"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1007\/s11101-006-9032-2","article-title":"Secondary metabolites from species of the biocontrol agent Trichoderma","volume":"7","author":"Reino","year":"2008","journal-title":"Phytochem. Rev."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1146\/annurev-arplant-042916-041132","article-title":"Defense priming: An adaptive part of induced resistance","volume":"68","author":"Baccelli","year":"2017","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"uhaf082","DOI":"10.1093\/hr\/uhaf082","article-title":"Salicylic acid and jasmonic acid in plant immunity","volume":"12","author":"Zhang","year":"2025","journal-title":"Hortic. Res."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1146\/annurev.phyto.42.040803.140421","article-title":"Systemic acquired resistance","volume":"42","author":"Durrant","year":"2002","journal-title":"Annu. Rev. Phytopathol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1016\/j.pbi.2007.04.020","article-title":"Networks of WRKY transcription factors in defense signaling","volume":"10","author":"Eulgem","year":"2007","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1016\/j.biocontrol.2013.07.005","article-title":"Diversity in plant systemic resistance induced by Trichoderma","volume":"67","author":"Nawrocka","year":"2013","journal-title":"Biol. Control"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1007\/s42452-026-08253-5","article-title":"Ecological versatility and biocontrol mechanisms of Trichoderma spp.: Toward sustainable agriculture","volume":"8","year":"2026","journal-title":"Discov. Appl. Sci."},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Yu, Y., Gui, Y., Li, Z., Jiang, C., Guo, J., and Niu, D. (2022). Induced systemic resistance for improving plant immunity by beneficial microbes. Plants, 11.","DOI":"10.3390\/plants11030386"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"818","DOI":"10.1016\/j.tplants.2016.07.009","article-title":"Recognizing plant defense priming","volume":"21","author":"Flors","year":"2016","journal-title":"Trends Plant Sci."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1094\/PDIS.2002.86.5.493","article-title":"Use of fungicides and biological controls in the suppression of Fusarium crown and root rot of asparagus under greenhouse and growth chamber conditions","volume":"86","author":"Reid","year":"2002","journal-title":"Plant Dis."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"100455","DOI":"10.1016\/j.stress.2024.100455","article-title":"The application of Trichoderma spp., an old but new useful fungus, in sustainable soil health intensification: A comprehensive strategy for addressing challenges","volume":"12","author":"Asghar","year":"2024","journal-title":"Plant Stress"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1007\/s12600-011-0151-y","article-title":"Trichoderma\u2014Not just for biocontrol anymore","volume":"39","author":"Harman","year":"2011","journal-title":"Phytoparasitica"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"21769","DOI":"10.1021\/acs.jafc.5c02157","article-title":"Using 13C15N labeling technology to reveal the fate of maize straw in soil organic matter and dissolved organic matter under different exogenous microorganism-mediated condition","volume":"73","author":"Zhang","year":"2025","journal-title":"J. Agric. Food Chem."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/S0065-2113(08)60948-7","article-title":"Growth promotion of plants inoculated with phosphate-solubilizing fungi","volume":"69","author":"Whitelaw","year":"1999","journal-title":"Adv. Agron."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"795","DOI":"10.1071\/CP20241","article-title":"Assessment of foliar-applied phosphorus fertiliser formulations to enhance phosphorus nutrition and grain production in wheat","volume":"71","author":"Mcbeath","year":"2020","journal-title":"Crop Pasture Sci."},{"key":"ref_71","first-page":"296","article-title":"Transcriptomic analysis of colonization by plant growth-promoting rhizobacterium Bacillus amyloliquefaciens SQR9 on gene expression of cucumber roots","volume":"44","author":"Ren","year":"2021","journal-title":"J. Nanjing Agric. Univ."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1007\/s00425-002-0921-3","article-title":"Molecular mechanisms of phosphate transport in plants","volume":"216","author":"Rausch","year":"2002","journal-title":"Planta"},{"key":"ref_73","first-page":"391","article-title":"Effect of five Trichoderma strains on growth and nutritional quality of Orchard grass","volume":"35","author":"Yuan","year":"2018","journal-title":"Pratacult. Sci."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"02027","DOI":"10.1051\/e3sconf\/202125102027","article-title":"Establishment of a rapid detection method for wheat crown rot","volume":"251","author":"Sui","year":"2021","journal-title":"E3S Web Conf."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1007\/s41348-016-0001-8","article-title":"Assessment of inoculation methods to identify resistance to Fusarium crown rot in wheat","volume":"123","author":"Poole","year":"2016","journal-title":"J. Plant Dis. Prot."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Yang, X., Pan, Y., Singh, P.K., He, X., Ren, Y., Zhao, L., Zhang, N., Cheng, S., and Chen, F. (2019). Investigation and genome-wide association study for Fusarium crown rot resistance in Chinese common wheat. BMC Plant Biol., 19.","DOI":"10.1186\/s12870-019-1758-2"},{"key":"ref_77","first-page":"431","article-title":"Identification and evaluation of wheat germplasm resistance to crown rot caused by Fusarium graminearum","volume":"3","author":"Zhang","year":"2009","journal-title":"J. Plant Genet. Resour."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1111\/j.1439-0434.2008.01425.x","article-title":"A simple method for the assessment of crown rot disease severity in wheat seedlings inoculated with Fusarium pseudograminearum","volume":"156","author":"Li","year":"2008","journal-title":"J. Phytopathol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/S0168-1656(84)90063-4","article-title":"Interspecific protoplast fusion of Trichoderma","volume":"1","author":"Toyama","year":"1984","journal-title":"J. Biotechnol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1080\/01621459.1931.10503208","article-title":"The test of significance for the Correlation Coefficient","volume":"26","author":"Pearson","year":"1931","journal-title":"J. Am. Stat. Assoc."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1198\/106186006X113430","article-title":"Sparse Principal Component Analysis","volume":"15","author":"Zou","year":"2006","journal-title":"J. Comput. Graph. Stat."},{"key":"ref_82","doi-asserted-by":"crossref","unstructured":"Li, J., Zhang, J., Li, H., Niu, H., Xu, Q., Jiao, Z., An, J., Jiang, Y., Li, Q., and Niu, J. (2019). The major factors causing the microspore abortion of genic male sterile mutant NWMS1 in wheat (Triticum aestivum L.). Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20246252"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"He, R., Ni, Y., J., L., Jiao, Z., Zhu, X., Jiang, Y., Li, Q., and Niu, J. (2018). Quantitative changes in the transcription of phytohormone-related genes: Some transcription factors are major causes of the wheat mutant dmc not tillering. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19051324"},{"key":"ref_84","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\u2212\u2206\u2206Ct method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"e11857","DOI":"10.7717\/peerj.11857","article-title":"Genome-wide association study of yield components in spring wheat collection harvested under two water regimes in Northern Kazakhstan","volume":"9","author":"Amalova","year":"2021","journal-title":"PeerJ"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1186\/s43170-023-00153-7","article-title":"Nitrogen uptake and remobilization from pre- and post-anthesis stages contribute towards grain yield and grain protein concentration in wheat grown in limited nitrogen conditions","volume":"4","author":"Sharma","year":"2023","journal-title":"CABI Agric. Biosci."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Alkahtani, J., Elshikh, M., Alwahibi, M., Muhammad, A., Ahmad, M., and Khalid, S. (2020). Phosphorus and zinc fertilization influence crop growth rates and total biomass of coarse vs. fine types rice cultivars. Agronomy, 10.","DOI":"10.3390\/agronomy10091356"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/19\/2957\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T12:05:50Z","timestamp":1790597150000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/19\/2957"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,28]]},"references-count":87,"journal-issue":{"issue":"19","published-online":{"date-parts":[[2026,10]]}},"alternative-id":["plants15192957"],"URL":"https:\/\/doi.org\/10.3390\/plants15192957","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,28]]}}}