{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T15:13:45Z","timestamp":1786115625059,"version":"build-2736575974"},"reference-count":97,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T00:00:00Z","timestamp":1785801600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"crossref","award":["32260303"],"award-info":[{"award-number":["32260303"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"crossref"}]},{"award":["32260303"],"award-info":[{"award-number":["32260303"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Major Scientific Research Project of Higher Education Institutions in Anhui Province","award":["2024AH040086"],"award-info":[{"award-number":["2024AH040086"]}]},{"name":"Anhui Province University Natural Science Research Foundation","award":["KJ2024AH051402"],"award-info":[{"award-number":["KJ2024AH051402"]}]},{"award":["KJ2024AH051402"],"award-info":[{"award-number":["KJ2024AH051402"]}],"id":[{"id":"https:\/\/ror.org\/05th9sq46","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Hubei Minzu University Scientific Research Startup Fund for Doctors","award":["BS25085"],"award-info":[{"award-number":["BS25085"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["J. Fungi","JoF"],"abstract":"<jats:p>Abiotic stresses increasingly threaten crop productivity, whereas reliance on chemical and resource-intensive interventions can compromise environmental sustainability. Existing literature identifies Trichoderma spp. as multifunctional biocontrol agents, biofertilizers, and microbial biostimulants capable of influencing plant growth, stress signaling, and rhizosphere processes; however, evidence remains fragmented across strains, crops, formulations, and stress conditions. This review aimed to integrate current knowledge on Trichoderma-mediated resilience to salinity, drought, heavy metals, temperature extremes, and emerging pollutants, while distinguishing experimentally validated mechanisms from statistical associations and conceptual inference. It evaluates constraints governing reproducibility from controlled studies to field deployment. The synthesis shows that selected crop\u2013strain systems improve root architecture, photosynthesis, antioxidant regulation, osmotic adjustment, nutrient acquisition, ion homeostasis, hormonal balance, and stress-responsive gene expression. Benefits arise through coordinated delivery, root colonization, metabolite and protein signaling, physiological reprogramming, and rhizosphere modulation. Nevertheless, microbiome co-occurrence patterns do not establish causal network repair, evidence for broad heat and cold protection remains limited, and biochar co-application should not be interpreted as a carrier formulation without direct validation. Future progress requires strain- and crop-specific screening, mechanistic gene and protein studies, standardized formulations, combined-stress experiments, multi-location field trials, biosafety evaluation, and farmer-level economic assessment to develop reliable precision microbial technologies.<\/jats:p>","DOI":"10.3390\/jof12080578","type":"journal-article","created":{"date-parts":[[2026,8,4]],"date-time":"2026-08-04T12:32:43Z","timestamp":1785846763000},"page":"578","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Trichoderma-Enabled Crop Resilience Under Abiotic Stress: From Field Delivery to Systems-Level Stress Reprogramming"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4681-6817","authenticated-orcid":false,"given":"Xueping","family":"Su","sequence":"first","affiliation":[{"name":"Hubei Key Laboratory of Biologic Resources Protection and Utilization, Hubei Minzu University, Enshi 445000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fangzhao","family":"Qin","sequence":"additional","affiliation":[{"name":"Hubei Key Laboratory of Biologic Resources Protection and Utilization, Hubei Minzu University, Enshi 445000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cheng","family":"Huang","sequence":"additional","affiliation":[{"name":"Hubei Key Laboratory of Biologic Resources Protection and Utilization, Hubei Minzu University, Enshi 445000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2523-8803","authenticated-orcid":false,"given":"Fasih Ullah","family":"Haider","sequence":"additional","affiliation":[{"name":"South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leiru","family":"Chen","sequence":"additional","affiliation":[{"name":"South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, China"},{"name":"Anhui Academy of Forestry, Hefei 230031, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,4]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Kopeck\u00e1, R., Kameniarov\u00e1, M., \u010cern\u00fd, M., Brzobohat\u00fd, B., and Nov\u00e1k, J. (2023). Abiotic Stress in Crop Production. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24076603"},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Jeyasri, R., Muthuramalingam, P., Satish, L., Pandian, S.K., Chen, J.T., Ahmar, S., Wang, X., Mora-Poblete, F., and Ramesh, M. (2021). An Overview of Abiotic Stress in Cereal Crops: Negative Impacts, Regulation, Biotechnology and Integrated Omics. Plants, 10.","DOI":"10.3390\/plants10071472"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"588","DOI":"10.1016\/j.tplants.2021.02.011","article-title":"Global Warming, Climate Change, and Environmental Pollution: Recipe for a Multifactorial Stress Combination Disaster","volume":"26","author":"Zandalinas","year":"2021","journal-title":"Trends Plant Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1146\/annurev.arplant.59.032607.092911","article-title":"Mechanisms of Salinity Tolerance","volume":"59","author":"Munns","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Wang, F., Sun, X., Wang, K., Long, B., Li, F., and Xie, D. (2025). Physiological and Multi-Omics Insights into Trichoderma harzianum Alleviating Aged Microplastic Stress in Nicotiana benthamiana. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms262311667"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1051\/agro:2008021","article-title":"Plant drought stress: Effects, mechanisms and management","volume":"29","author":"Farooq","year":"2009","journal-title":"Agron. Sustain. Dev."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1016\/j.plaphy.2010.08.016","article-title":"Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants","volume":"48","author":"Gill","year":"2010","journal-title":"Plant Physiol. Biochem."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Hasanuzzaman, M., Nahar, K., and Fujita, M. (2013). Plant Response to Salt Stress and Role of Exogenous Protectants to Mitigate Salt-Induced Damages. Ecophysiology and Responses of Plants Under Salt Stress, Springer.","DOI":"10.1007\/978-1-4614-4747-4_2"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/j.envexpbot.2007.05.011","article-title":"Heat tolerance in plants: An overview","volume":"61","author":"Wahid","year":"2007","journal-title":"Environ. Exp. Bot."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"111887","DOI":"10.1016\/j.ecoenv.2020.111887","article-title":"Cadmium toxicity in plants: Impacts and remediation strategies","volume":"211","author":"Haider","year":"2021","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"S\u00e1nchez-Berm\u00fadez, M., Del Pozo, J.C., and Pernas, M. (2022). Effects of Combined Abiotic Stresses Related to Climate Change on Root Growth in Crops. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.918537"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1093\/jxb\/err269","article-title":"Any trait or trait-related allele can confer drought tolerance: Just design the right drought scenario","volume":"63","author":"Tardieu","year":"2012","journal-title":"J. Exp. Bot."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1038\/nrg3901","article-title":"Genetic mechanisms of abiotic stress tolerance that translate to crop yield stability","volume":"16","author":"Mickelbart","year":"2015","journal-title":"Nat. Rev. Genet."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Seleiman, M.F., Al-Suhaibani, N., Ali, N., Akmal, M., Alotaibi, M., Refay, Y., Dindaroglu, T., Abdul-Wajid, H.H., and Battaglia, M.L. (2021). Drought Stress Impacts on Plants and Different Approaches to Alleviate Its Adverse Effects. Plants, 10.","DOI":"10.3390\/plants10020259"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"105078","DOI":"10.1016\/j.envint.2019.105078","article-title":"Soil and the intensification of agriculture for global food security","volume":"132","author":"Kopittke","year":"2019","journal-title":"Environ. Int."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Godoy, F., Olivos-Hern\u00e1ndez, K., Stange, C., and Handford, M. (2021). Abiotic Stress in Crop Species: Improving Tolerance by Applying Plant Metabolites. Plants, 10.","DOI":"10.3390\/plants10020186"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1386","DOI":"10.1016\/j.enceco.2025.06.009","article-title":"Synergistic regulation of Cd stress tolerance in Brassica juncea: Metabolic reprogramming and nutrient-Cd co-transport under Trichoderma harzianum and polyaspartic acid","volume":"7","author":"Yao","year":"2025","journal-title":"Environ. Chem. Ecotoxicol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"127621","DOI":"10.1016\/j.micres.2024.127621","article-title":"Mechanisms for plant growth promotion activated by Trichoderma in natural and managed terrestrial ecosystems","volume":"281","author":"Schmoll","year":"2024","journal-title":"Microbiol. Res."},{"key":"ref_19","first-page":"2261","article-title":"The Mechanisms of Trichoderma Species to Reduce Drought and Salinity Stress in Plants","volume":"92","author":"Boorboori","year":"2023","journal-title":"Phyton-Int. J. Exp. Bot."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Geng, Y., Chen, S., Lv, P., Li, Y., Li, J., Jiang, F., Wu, Z., Shen, Q., and Zhou, R. (2025). Positive Role of Trichoderma harzianum in Increasing Plant Tolerance to Abiotic Stresses: A Review. Antioxidants, 14.","DOI":"10.3390\/antiox14070807"},{"key":"ref_21","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_22","doi-asserted-by":"crossref","unstructured":"Yakhin, O.I., Lubyanov, A.A., Yakhin, I.A., and Brown, P.H. (2017). Biostimulants in Plant Science: A Global Perspective. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.02049"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1111\/j.1469-8137.2010.03614.x","article-title":"Multifunctional fungal plant symbionts: New tools to enhance plant growth and productivity","volume":"189","author":"Harman","year":"2011","journal-title":"New Phytol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1038\/nrmicro797","article-title":"Trichoderma species\u2014Opportunistic, avirulent plant symbionts","volume":"2","author":"Harman","year":"2004","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"e41114","DOI":"10.1016\/j.heliyon.2024.e41114","article-title":"Application of Trichoderma harzianum enhances salt tolerance and yield of Indian mustard through increasing antioxidant enzyme activity","volume":"11","author":"Saha","year":"2025","journal-title":"Heliyon"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1264","DOI":"10.1094\/MPMI-09-11-0240","article-title":"Trichoderma harzianum enhances antioxidant defense of tomato seedlings and resistance to water deficit","volume":"25","author":"Mastouri","year":"2012","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1016\/j.enceco.2025.06.001","article-title":"Synergistic effects of Trichoderma harzianum and polyaspartic acid enhance cadmium phytoremediation via core microbiota and network stability in Brassica juncea","volume":"7","author":"Yao","year":"2025","journal-title":"Environ. Chem. Ecotoxicol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1007\/s11274-024-03965-5","article-title":"Recent advances in the use of Trichoderma-containing multicomponent microbial inoculants for pathogen control and plant growth promotion","volume":"40","author":"Kredics","year":"2024","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Bejarano, A., and Puopolo, G. (2020). Bioformulation of Microbial Biocontrol Agents for a Sustainable Agriculture. How Research Can Stimulate the Development of Commercial Biological Control Against Plant Diseases, Springer.","DOI":"10.1007\/978-3-030-53238-3_16"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Teixid\u00f3, N., Segarra, G., Casals, C., Usall, J., and Torres, R. (2020). Formulations to Improve Biocontrol Products Shelf-Life and\/or Ecosystem Adaptation. How Research Can Stimulate the Development of Commercial Biological Control Against Plant Diseases, Springer.","DOI":"10.1007\/978-3-030-53238-3_15"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1146\/annurev-phyto-082712-102353","article-title":"Trichoderma research in the genome era","volume":"51","author":"Mukherjee","year":"2013","journal-title":"Annu. Rev. Phytopathol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"111193","DOI":"10.1016\/j.plaphy.2026.111193","article-title":"Trichoderma in multitrophic plant-microbe interactions: A pan-genome guided roadmap for resilient physiology and sustainable bio-economy","volume":"232","author":"Mondal","year":"2026","journal-title":"Plant Physiol. Biochem."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Khan, R.A., Najeeb, S., Hussain, S., Xie, B., and Li, Y. (2020). Bioactive Secondary Metabolites from Trichoderma spp. Against Phytopathogenic Fungi. Microorganisms , 8.","DOI":"10.3390\/microorganisms8060817"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"127","DOI":"10.2174\/1874437001408010127","article-title":"Trichoderma secondary metabolites active on plants and fungal pathogens","volume":"8","author":"Vinale","year":"2014","journal-title":"Open Mycol. J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1111\/nph.16530","article-title":"Sniffing fungi-phenotyping of volatile chemical diversity in Trichoderma species","volume":"227","author":"Guo","year":"2020","journal-title":"New Phytol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1496","DOI":"10.1111\/nph.13725","article-title":"The volatile 6-pentyl-2H-pyran-2-one from Trichoderma atroviride regulates Arabidopsis thaliana root morphogenesis via auxin signaling and ETHYLENE INSENSITIVE 2 functioning","volume":"209","year":"2016","journal-title":"New Phytol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.fgb.2017.01.002","article-title":"HFB7-A novel orphan hydrophobin of the Harzianum and Virens clades of Trichoderma, is involved in response to biotic and abiotic stresses","volume":"102","author":"Przylucka","year":"2017","journal-title":"Fungal Genet. Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1111\/j.1364-3703.2006.00335.x","article-title":"TasHyd1, a new hydrophobin gene from the biocontrol agent Trichoderma asperellum, is involved in plant root colonization","volume":"7","author":"Viterbo","year":"2006","journal-title":"Mol. Plant Pathol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"2926","DOI":"10.1128\/AEM.65.7.2926-2933.1999","article-title":"Solubilization of phosphates and micronutrients by the plant-growth-promoting and biocontrol fungus Trichoderma harzianum Rifai 1295-22","volume":"65","author":"Altomare","year":"1999","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"105100","DOI":"10.1016\/j.biocontrol.2022.105100","article-title":"Combined use of Trichoderma and beneficial bacteria (mainly Bacillus and Pseudomonas): Development of microbial synergistic bio-inoculants in sustainable agriculture","volume":"176","author":"Poveda","year":"2022","journal-title":"Biol. Control"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"100360","DOI":"10.1016\/j.microb.2025.100360","article-title":"Trichoderma-bacterial network: A balance inter-kingdom interaction for agricultural relevance","volume":"7","author":"Othman","year":"2025","journal-title":"Microbe"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1111\/1462-2920.15878","article-title":"Dual inoculation of dark septate endophytes and Trichoderma viride drives plant performance and rhizosphere microbiome adaptations of Astragalus mongholicus to drought","volume":"24","author":"He","year":"2022","journal-title":"Environ. Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1061","DOI":"10.1128\/AEM.65.3.1061-1070.1999","article-title":"Induction of defense responses in cucumber plants (Cucumis sativus L.) BY Biocontrol Agent Trichoderma harzianum","volume":"65","author":"Yedidia","year":"1999","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/j.funeco.2012.09.005","article-title":"Arabidopsis thaliana as a model system for testing the effect of Trichoderma volatile organic compounds","volume":"6","author":"Hung","year":"2013","journal-title":"Fungal Ecol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1579","DOI":"10.1104\/pp.108.130369","article-title":"Trichoderma virens, a Plant Beneficial Fungus, Enhances Biomass Production and Promotes Lateral Root Growth through an Auxin-Dependent Mechanism in Arabidopsis","volume":"149","year":"2009","journal-title":"Plant Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1968","DOI":"10.1016\/j.soilbio.2007.02.015","article-title":"Growth stimulation and fruit yield improvement of greenhouse tomato plants by inoculation with Pseudomonas putida or Trichoderma atroviride: Possible role of indole acetic acid (IAA)","volume":"39","author":"Gravel","year":"2007","journal-title":"Soil Biol. Biochem."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1111\/j.1574-6968.2010.01910.x","article-title":"Characterization of ACC deaminase from the biocontrol and plant growth-promoting agent Trichoderma asperellum T203","volume":"305","author":"Viterbo","year":"2010","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1007\/s11104-017-3445-0","article-title":"Modulating rhizosphere colonisation, plant growth, soil nutrient availability and plant defense enzyme activity through Trichoderma viride-Azotobacter chroococcum biofilm inoculation in chickpea","volume":"421","author":"Velmourougane","year":"2017","journal-title":"Plant Soil"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Guzm\u00e1n-Guzm\u00e1n, P., Etesami, H., and Santoyo, G. (2025). Trichoderma: A multifunctional agent in plant health and microbiome interactions. BMC Microbiol., 25.","DOI":"10.1186\/s12866-025-04158-2"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"478","DOI":"10.1016\/j.tplants.2012.04.001","article-title":"The rhizosphere microbiome and plant health","volume":"17","author":"Berendsen","year":"2012","journal-title":"Trends Plant Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1007\/s11104-014-2326-z","article-title":"Colonization of Trichoderma harzianum strain SQR-T037 on tomato roots and its relationship to plant growth, nutrient availability and soil microflora","volume":"388","author":"Cai","year":"2015","journal-title":"Plant Soil"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1099\/mic.0.052274-0","article-title":"Plant-beneficial effects of Trichoderma and of its genes","volume":"158","author":"Hermosa","year":"2012","journal-title":"Microbiology"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Brotman, Y., Landau, U., Cuadros-Inostroza, \u00c1., Tohge, T., Fernie, A.R., Chet, I., Viterbo, A., and Willmitzer, L. (2013). Trichoderma-plant root colonization: Escaping early plant defense responses and activation of the antioxidant machinery for saline stress tolerance. PLoS Pathog., 9.","DOI":"10.1371\/annotation\/8b818c15-3fe0-4e56-9be2-e44fd1ed3fae"},{"key":"ref_54","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_55","doi-asserted-by":"crossref","first-page":"7019","DOI":"10.1007\/s00344-024-11474-w","article-title":"Trichoderma harzianum Protects the Arabidopsis Salt Overly Sensitive 1 Mutant Against Salt Stress","volume":"44","author":"Gandhi","year":"2025","journal-title":"J. Plant Growth Regul."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Zhao, M., Wang, P., Liu, X., and Jin, L. (2026). The Effects of Trichoderma harzianum Inoculation on the Growth, Nutrient Absorption, and Expressions of Stress-Responsive Genes of Citrus Under Salt Stress. Horticulturae, 12.","DOI":"10.3390\/horticulturae12020233"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"100370","DOI":"10.1016\/j.stress.2024.100370","article-title":"The role of Trichoderma koningii and Trichoderma harzianum in mitigating the combined stresses motivated by Sclerotinia sclerotiorum and salinity in common bean (Phaseolus vulgaris)","volume":"11","author":"Abdelrhim","year":"2024","journal-title":"Plant Stress"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Ahmad, P., Hashem, A., Abd-Allah, E.F., Alqarawi, A.A., John, R., Egamberdieva, D., and Gucel, S. (2015). Role of Trichoderma harzianum in mitigating NaCl stress in Indian mustard (Brassica juncea L.) through the antioxidative defense system. Front. Plant Sci., 6.","DOI":"10.3389\/fpls.2015.00868"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1186\/s40529-023-00368-x","article-title":"Alleviation of the adverse effects of NaCl stress on tomato seedlings (Solanum lycopersicum L.) by Trichoderma viride through the antioxidative defense system","volume":"64","author":"Metwally","year":"2023","journal-title":"Bot. Stud."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Bashyal, B.M., Parmar, P., Zaidi, N.W., and Aggarwal, R. (2021). Molecular Programming of Drought-Challenged Trichoderma harzianum-Bioprimed Rice (Oryza sativa L.). Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.655165"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.plaphy.2017.10.012","article-title":"Overexpression of an aquaglyceroporin gene from Trichoderma harzianum improves water-use efficiency and drought tolerance in Nicotiana tabacum","volume":"121","author":"Vieira","year":"2017","journal-title":"Plant Physiol. Biochem."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1007\/s00425-020-03404-3","article-title":"Application of Trichoderma asperellum T34 on maize (Zea mays) seeds protects against drought stress","volume":"252","author":"Vicente","year":"2020","journal-title":"Planta"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Scudeletti, D., Crusciol, C.A., Bossolani, J.W., Moretti, L.G., Momesso, L., Servaz Tuba\u00f1a, B., De Castro, S.G., De Oliveira, E.F., and Hungria, M. (2021). Trichoderma asperellum Inoculation as a Tool for Attenuating Drought Stress in Sugarcane. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.645542"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Cs\u00f3t\u00f3, A., T\u00f3th, G., Riczu, P., Zabi\u00e1k, A., Tarj\u00e1nyi, V., Fekete, E., Karaffa, L., and S\u00e1ndor, E. (2024). Foliar Spraying with Endophytic Trichoderma Biostimulant Increases Drought Resilience of Maize and Sunflower. Agriculture, 14.","DOI":"10.3390\/agriculture14122360"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1751","DOI":"10.1016\/S2095-3119(17)61695-2","article-title":"Increased resistance of drought by Trichoderma harzianum fungal treatment correlates with increased secondary metabolites and proline content","volume":"16","author":"Mona","year":"2017","journal-title":"J. Integr. Agric."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1111\/aab.12454","article-title":"The influence of Trichoderma brevicompactum treatment and drought on physiological parameters, abscisic acid content and signalling pathway marker gene expression in leaves and roots of tomato","volume":"173","year":"2018","journal-title":"Ann. Appl. Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1007\/s10725-025-01423-2","article-title":"Physiological and biochemical ameliorative effects of biochar, Trichoderma harzianum, and combined applications in Capsicum annuum L. under water stress","volume":"106","author":"Kocaman","year":"2026","journal-title":"Plant Growth Regul."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1186\/s13213-025-01800-2","article-title":"Molecular and genetic adaptations of heat-treated Trichoderma harzianum for enhanced stress resilience and biocontrol efficiency","volume":"75","author":"Ahmed","year":"2025","journal-title":"Ann. Microbiol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"2005","DOI":"10.1080\/03650340.2021.1949709","article-title":"Integrated usage of Trichoderma harzianum and biochar to ameliorate salt stress on spinach plants","volume":"68","author":"Sofy","year":"2022","journal-title":"Arch. Agron. Soil Sci."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1111\/jam.14368","article-title":"Endophytic strains of Trichoderma increase plants\u2019 photosynthetic capability","volume":"130","author":"Harman","year":"2021","journal-title":"J. Appl. Microbiol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Rawal, R., Scheerens, J.C., Fenstemaker, S.M., Francis, D.M., Miller, S.A., and Benitez, M.S. (2022). Novel Trichoderma Isolates Alleviate Water Deficit Stress in Susceptible Tomato Genotypes. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.869090"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"804","DOI":"10.1007\/s10886-014-0478-1","article-title":"Phytohormone Profiles Induced by Trichoderma Isolates Correspond with Their Biocontrol and Plant Growth-Promoting Activity on Melon Plants","volume":"40","author":"Pascual","year":"2014","journal-title":"J. Chem. Ecol."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Zhang, S., Xu, B., and Gan, Y. (2019). Seed Treatment with Trichoderma longibrachiatum T6 Promotes Wheat Seedling Growth under NaCl Stress Through Activating the Enzymatic and Nonenzymatic Antioxidant Defense Systems. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20153729"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Gupta, S.V., Smith, P.M., Natera, S.H., and Roessner, U. (2022). Biochemical Changes in Two Barley Genotypes Inoculated With a Beneficial Fungus Trichoderma harzianum Rifai T-22 Grown in Saline Soil. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.908853"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1104\/pp.108.116293","article-title":"Role of Swollenin, an Expansin-Like Protein from Trichoderma, in Plant Root Colonization","volume":"147","author":"Brotman","year":"2008","journal-title":"Plant Physiol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"875","DOI":"10.1104\/pp.107.103689","article-title":"A Proteinaceous Elicitor Sm1 from the Beneficial Fungus Trichoderma virens Is Required for Induced Systemic Resistance in Maize","volume":"145","author":"Vargas","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Poveda, J., Hermosa, R., Monte, E., and Nicol\u00e1s, C. (2019). The Trichoderma harzianum Kelch Protein ThKEL1 Plays a Key Role in Root Colonization and the Induction of Systemic Defense in Brassicaceae Plants. Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.01478"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Druzhinina, I.S., Chenthamara, K., Zhang, J., Atanasova, L., Yang, D., Miao, Y., Rahimi, M.J., Grujic, M., Cai, F., and Pourmehdi, S. (2018). Massive lateral transfer of genes encoding plant cell wall-degrading enzymes to the mycoparasitic fungus Trichoderma from its plant-associated hosts. PLoS Genet., 14.","DOI":"10.1371\/journal.pgen.1007322"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1038\/s41579-020-0412-1","article-title":"Plant-microbiome interactions: From community assembly to plant health","volume":"18","author":"Trivedi","year":"2020","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"963401","DOI":"10.6064\/2012\/963401","article-title":"Plant Growth-Promoting Bacteria: Mechanisms and Applications","volume":"2012","author":"Glick","year":"2012","journal-title":"Scientifica"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1812","DOI":"10.1016\/j.soilbio.2011.04.022","article-title":"Biochar effects on soil biota\u2014A review","volume":"43","author":"Lehmann","year":"2011","journal-title":"Soil Biol. Biochem."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/S0065-2113(10)05002-9","article-title":"A review of biochar and its use and function in soil","volume":"105","author":"Sohi","year":"2010","journal-title":"Adv. Agron."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1038\/s41579-018-0024-1","article-title":"Keystone taxa as drivers of microbiome structure and functioning","volume":"16","author":"Banerjee","year":"2018","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"2426","DOI":"10.1038\/ismej.2017.91","article-title":"Cohesion: A method for quantifying the connectivity of microbial communities","volume":"11","author":"Herren","year":"2017","journal-title":"ISME J."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"5729","DOI":"10.1007\/s00253-016-7590-9","article-title":"Microbial inoculation of seed for improved crop performance: Issues and opportunities","volume":"100","year":"2016","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11104-013-1956-x","article-title":"Advances in plant growth-promoting bacterial inoculant technology: Formulations and practical perspectives (1998\u20132013)","volume":"378","author":"Bashan","year":"2014","journal-title":"Plant Soil"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"863240","DOI":"10.1155\/2013\/863240","article-title":"Microbial inoculants and their impact on soil microbial communities: A review","volume":"2013","author":"Trabelsi","year":"2013","journal-title":"BioMed Res. Int."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1038\/s41477-018-0139-4","article-title":"Core microbiomes for sustainable agroecosystems","volume":"4","author":"Toju","year":"2018","journal-title":"Nat. Plants"},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Backer, R., Rokem, J.S., Ilangumaran, G., Lamont, J., Praslickova, D., Ricci, E., Subramanian, S., and Smith, D.L. (2018). Plant Growth-Promoting Rhizobacteria: Context, Mechanisms of Action, and Roadmap to Commercialization of Biostimulants for Sustainable Agriculture. Front. Plant Sci., 9.","DOI":"10.3389\/fpls.2018.01473"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1007\/s10526-012-9498-3","article-title":"Regulating biocontrol agents: A historical perspective and a critical examination comparing microbial and macrobial agents","volume":"58","author":"Sundh","year":"2013","journal-title":"BioControl"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.jare.2019.03.004","article-title":"A review on the plant microbiome: Ecology, functions, and emerging trends in microbial application","volume":"19","author":"Compant","year":"2019","journal-title":"J. Adv. Res."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.pbi.2017.04.018","article-title":"Understanding and exploiting plant beneficial microbes","volume":"38","author":"Finkel","year":"2017","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Busby, P.E., Soman, C., Wagner, M.R., Friesen, M.L., Kremer, J., Bennett, A., Morsy, M., Eisen, J.A., Leach, J.E., and Dangl, J.L. (2017). Research priorities for harnessing plant microbiomes in sustainable agriculture. PLoS Biol., 15.","DOI":"10.1371\/journal.pbio.2001793"},{"key":"ref_94","first-page":"100094","article-title":"Biofertilizers: An ecofriendly technology for nutrient recycling and environmental sustainability","volume":"3","author":"Kumar","year":"2022","journal-title":"Curr. Res. Microb. Sci."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.scienta.2015.09.021","article-title":"Plant biostimulants: Definition, concept, main categories and regulation","volume":"196","year":"2015","journal-title":"Sci. Hortic."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"6599","DOI":"10.1007\/s00253-014-5828-y","article-title":"A contribution to set a legal framework for biofertilisers","volume":"98","author":"Malusa","year":"2014","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_97","doi-asserted-by":"crossref","unstructured":"Mitter, E.K., Tosi, M., Obreg\u00f3n, D., Dunfield, K.E., and Germida, J.J. (2021). Rethinking Crop Nutrition in Times of Modern Microbiology: Innovative Biofertilizer Technologies. Front. Sustain. 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