{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T05:46:02Z","timestamp":1785735962426,"version":"3.56.0"},"reference-count":69,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T00:00:00Z","timestamp":1785715200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>Fungal pathogens are a vital component of microbial communities and can substantially reduce plant productivity. In grasslands, pathogens and their host plants frequently coexist with large grazing herbivores. However, it remains unclear whether and how grazing alters pathogen transmission and prevalence, particularly when integrating soil and foliar pathogens. Here, we conducted an 8-year field experiment in alpine grasslands on the Qinghai-Tibetan Plateau to examine how soil pathogens and foliar diseases respond to grazing intensity, and to assess the relationship between them. Our results showed that light grazing significantly increased the incidence and severity of plant community foliar diseases, while moderate and heavy grazing significantly reduced them. Similarly, light grazing significantly increased the relative abundance of soil pathogens. Notably, we found a strong positive relationship between soil pathogen abundance and foliar disease occurrence. Mechanistically, light grazing indirectly exacerbated foliar diseases through increases in soil pathogen abundance, whereas moderate and heavy grazing directly suppressed them. Overall, our findings highlight the distinct effects of different grazing intensities on soil pathogens and foliar diseases, and further provide experimental evidence that soil pathogens may play an important role in foliar disease dynamics in grassland ecosystems.<\/jats:p>","DOI":"10.3389\/fpls.2026.1900475","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T05:30:30Z","timestamp":1785735030000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Effects of grazing intensity on plant community fungal diseases in an alpine grassland"],"prefix":"10.3389","volume":"17","author":[{"given":"Zhen","family":"Tian","sequence":"first","affiliation":[{"name":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems; Center for Grassland Microbiome; College of Pastoral Agricultural Science and Technology, Lanzhou University","place":["Lanzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoxia","family":"Yang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Plateau Ecology and Agriculture, Qinghai Academy of Animal and Veterinary Science, Qinghai University","place":["Xining, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Quanmin","family":"Dong","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Plateau Ecology and Agriculture, Qinghai Academy of Animal and Veterinary Science, Qinghai University","place":["Xining, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tao","family":"Chen","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems; Center for Grassland Microbiome; College of Pastoral Agricultural Science and Technology, Lanzhou University","place":["Lanzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,8,3]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"2572","DOI":"10.1890\/09-0859","article-title":"Foliar fungal pathogens and grassland biodiversity","volume":"91","author":"Allan","year":"2010","journal-title":"Ecology"},{"key":"B2","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1126\/science.abo2380","article-title":"Grassland soil carbon sequestration: Current understanding, challenges, and solutions","volume":"377","author":"Bai","year":"2022","journal-title":"Science"},{"key":"B3","doi-asserted-by":"publisher","first-page":"2258","DOI":"10.1890\/02-0274","article-title":"Herbivore\u2010mediated linkages between aboveground and belowground communities","volume":"84","author":"Bardgett","year":"2003","journal-title":"Ecology"},{"key":"B4","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1016\/j.still.2010.05.010","article-title":"Evaluation of soil compaction effects on soil biota and soil biological processes in soils","volume":"109","author":"Beylich","year":"2010","journal-title":"Soil Tillage Res."},{"key":"B5","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1073\/pnas.0808778106","article-title":"Consumers indirectly increase infection risk in grassland food webs","volume":"106","author":"Borer","year":"2009","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B6","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1038\/nmeth.3869","article-title":"DADA2: High-resolution sample inference from Illumina amplicon data","volume":"13","author":"Callahan","year":"2016","journal-title":"Nat. Methods"},{"key":"B7","doi-asserted-by":"publisher","first-page":"1349","DOI":"10.1111\/ele.13537","article-title":"Sick plants in grassland communities: a growth\u2010defense trade\u2010off is the main driver of fungal pathogen abundance","volume":"23","author":"Cappelli","year":"2020","journal-title":"Ecol. Lett."},{"key":"B8","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1038\/s41586-024-08419-4","article-title":"Fungal impacts on Earth\u2019s ecosystems","volume":"638","author":"Case","year":"2025","journal-title":"Nature"},{"key":"B9","doi-asserted-by":"publisher","first-page":"781","DOI":"10.1111\/j.1365-2745.2009.01508.x","article-title":"Grazer facilitation of fungal infection and the control of plant growth in south-western Atlantic salt marshes","volume":"97","author":"Daleo","year":"2009","journal-title":"J. Ecol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1038\/s41558-020-0759-3","article-title":"The proportion of soil-borne pathogens increases with warming at the global scale","volume":"10","author":"Delgado-Baquerizo","year":"2020","journal-title":"Nat. Clim. Change"},{"key":"B11","doi-asserted-by":"publisher","first-page":"e01337","DOI":"10.1128\/msystems.01337-21","article-title":"The proportion of soil-borne fungal pathogens increases with elevated organic carbon in agricultural soils","volume":"7","author":"Du","year":"2022","journal-title":"Msystems"},{"key":"B12","doi-asserted-by":"publisher","first-page":"1827","DOI":"10.1111\/1365-2435.14346","article-title":"Drought neutralizes positive effects of long\u2010term grazing on grassland productivity through altering plant\u2013soil interactions","volume":"37","author":"Duan","year":"2023","journal-title":"Funct. Ecol."},{"key":"B13","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1111\/j.1365-3059.2010.02402.x","article-title":"Influence of atmospheric and climatic change on plant\u2013pathogen interactions","volume":"60","author":"Eastburn","year":"2011","journal-title":"Plant Pathol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1890\/08-2338.1","article-title":"Plant diversity effects on soil microorganisms support the singular hypothesis","volume":"91","author":"Eisenhauer","year":"2010","journal-title":"Ecology"},{"key":"B15","doi-asserted-by":"publisher","first-page":"528","DOI":"10.1016\/j.tplants.2021.12.010","article-title":"Plant defense strategies against attack by multiple herbivores","volume":"27","author":"Fern\u00e1ndez de Bobadilla","year":"2022","journal-title":"Trends Plant Sci."},{"key":"B16","doi-asserted-by":"publisher","first-page":"186","DOI":"10.1038\/nature10947","article-title":"Emerging fungal threats to animal, plant and ecosystem health","volume":"484","author":"Fisher","year":"2012","journal-title":"Nature"},{"key":"B17","doi-asserted-by":"publisher","first-page":"344","DOI":"10.1038\/242344a0","article-title":"Competitive exclusion in herbaceous vegetation","volume":"242","author":"Grime","year":"1973","journal-title":"Nature"},{"key":"B18","doi-asserted-by":"publisher","first-page":"1992","DOI":"10.1111\/1365-2745.14370","article-title":"Ungulate herbivores promote beta diversity and drive stochastic plant community assembly by selective defoliation and trampling: From a four\u2010year simulation experiment","volume":"112","author":"Guo","year":"2024","journal-title":"J. Ecol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"2158","DOI":"10.1126\/science.1063699","article-title":"Climate warming and disease risks for terrestrial and marine biota","volume":"296","author":"Harvell","year":"2002","journal-title":"Science"},{"key":"B20","doi-asserted-by":"publisher","first-page":"757","DOI":"10.1111\/oik.04893","article-title":"Effects of grazing intensity on plant richness and diversity: A meta\u2010analysis","volume":"127","author":"Herrero\u2010J\u00e1uregui","year":"2018","journal-title":"Oikos"},{"key":"B21","doi-asserted-by":"publisher","first-page":"2738","DOI":"10.1038\/s41467-018-05122-7","article-title":"Root exudate metabolites drive plant-soil feedbacks on growth and defense by shaping the rhizosphere microbiota","volume":"9","author":"Hu","year":"2018","journal-title":"Nat. Commun."},{"key":"B22","doi-asserted-by":"publisher","first-page":"1001","DOI":"10.1038\/s41477-025-01987-x","article-title":"Herbivory-induced green leaf volatiles increase plant performance through jasmonate-dependent plant\u2013soil feedbacks","volume":"11","author":"Hu","year":"2025","journal-title":"Nat. Plants"},{"key":"B23","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1146\/annurev.py.12.090174.001035","article-title":"Nitrogen form and plant disease","volume":"12","author":"Huber","year":"1974","journal-title":"Annu. Rev. Phytopathol."},{"key":"B24","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1146\/annurev.py.30.090192.003005","article-title":"Modeling leaf wetness in relation to plant disease epidemiology","volume":"30","author":"Huber","year":"1992","journal-title":"Annu. Rev. Phytopathol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"1119","DOI":"10.1111\/ele.12479","article-title":"Frontiers in research on biodiversity and disease","volume":"18","author":"Johnson","year":"2015","journal-title":"Ecol. Lett."},{"key":"B26","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1111\/j.1461-0248.2006.00885.x","article-title":"Effects of species diversity on disease risk","volume":"9","author":"Keesing","year":"2006","journal-title":"Ecol. Lett."},{"key":"B27","doi-asserted-by":"publisher","first-page":"6707","DOI":"10.1038\/ncomms7707","article-title":"Plant diversity increases soil microbial activity and soil carbon storage","volume":"6","author":"Lange","year":"2015","journal-title":"Nat. Commun."},{"key":"B28","doi-asserted-by":"publisher","first-page":"597","DOI":"10.1111\/j.1365-2745.2011.01940.x","article-title":"Plant diversity improves protection against soil\u2010borne pathogens by fostering antagonistic bacterial communities","volume":"100","author":"Latz","year":"2012","journal-title":"J. Ecol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"3484","DOI":"10.1038\/s41467-021-23605-y","article-title":"Nitrogen and phosphorus fertilization consistently favor pathogenic over mutualistic fungi in grassland soils","volume":"12","author":"Lekberg","year":"2021","journal-title":"Nat. Commun."},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1016\/j.agee.2025.109516","article-title":"The global effects of grazing on grassland soil nitrogen retention","volume":"383","author":"Li","year":"2025","journal-title":"Agric. Ecosyst. Environ."},{"key":"B31","doi-asserted-by":"publisher","first-page":"6482","DOI":"10.1038\/s41467-023-42142-4","article-title":"Global diversity and biogeography of potential phytopathogenic fungi in a changing world","volume":"14","author":"Li","year":"2023","journal-title":"Nat. Commun."},{"key":"B32","doi-asserted-by":"publisher","first-page":"791","DOI":"10.1111\/1365-2664.14093","article-title":"Long\u2010term heavy grazing increases community\u2010level foliar fungal diseases by shifting plant composition","volume":"59","author":"Liu","year":"2022","journal-title":"J. Appl. Ecol."},{"key":"B33","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1111\/nph.17324","article-title":"Contrasting effects of mammal grazing on foliar fungal diseases: patterns and potential mechanisms","volume":"232","author":"Liu","year":"2021","journal-title":"New Phytol."},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2023.1117372","article-title":"Herbivore assemblages affect soil microbial communities by altering root biomass and available nutrients in an alpine meadow","volume":"14","author":"Liu","year":"2023","journal-title":"Front. Plant Sci."},{"key":"B35","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1111\/nph.17797","article-title":"Environmental and plant community drivers of plant pathogen composition and richness","volume":"233","author":"Makiola","year":"2022","journal-title":"New Phytol."},{"key":"B36","doi-asserted-by":"publisher","first-page":"1347","DOI":"10.1038\/s41477-021-00999-7","article-title":"Plant defence to sequential attack is adapted to prevalent herbivores","volume":"7","author":"Mertens","year":"2021","journal-title":"Nat. Plants"},{"key":"B37","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1086\/284839","article-title":"A generalized model of the effects of grazing by large herbivores on grassland community structure","volume":"132","author":"Milchunas","year":"1988","journal-title":"Am. Nat."},{"key":"B38","doi-asserted-by":"publisher","first-page":"897","DOI":"10.1007\/s00442-022-05223-7","article-title":"Plant community-mediated effects of grazing on plant diseases","volume":"199","author":"Mipam","year":"2022","journal-title":"Oecologia"},{"key":"B39","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1046\/j.1365-2486.2003.00602.x","article-title":"Effects of elevated CO2, nitrogen deposition, and decreased species diversity on foliar fungal plant disease","volume":"9","author":"Mitchell","year":"2003","journal-title":"Glob. Change Biol."},{"key":"B40","doi-asserted-by":"publisher","first-page":"2353","DOI":"10.1111\/1365-2745.12987","article-title":"Interactions between plant defence signalling pathways: evidence from bioassays with insect herbivores and plant pathogens","volume":"106","author":"Moreira","year":"2018","journal-title":"J. Ecol."},{"key":"B41","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/j.funeco.2015.06.006","article-title":"FUNGuild: an open annotation tool for parsing fungal community datasets by ecological guild","volume":"20","author":"Nguyen","year":"2016","journal-title":"Fungal Ecol."},{"key":"B42","doi-asserted-by":"publisher","first-page":"542","DOI":"10.1038\/nature14372","article-title":"Phylogenetic structure and host abundance drive disease pressure in communities","volume":"520","author":"Parker","year":"2015","journal-title":"Nature"},{"key":"B43","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1111\/j.1365-3059.1995.tb02773.x","article-title":"Fusarium ear blight (scab) in small grain cereals\u2014a review","volume":"44","author":"Parry","year":"1995","journal-title":"Plant Pathol."},{"key":"B44","doi-asserted-by":"publisher","first-page":"766","DOI":"10.1111\/geb.13662","article-title":"The climate\u2010driven distribution and response to global change of soil\u2010borne pathogens in agroecosystems","volume":"32","author":"Peng","year":"2023","journal-title":"Glob. Ecol. Biogeogr."},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.70345","article-title":"Grazing modulates the multiscale spatial structure of dryland vegetation","volume":"31","author":"Pichon","year":"2025","journal-title":"Glob. Change Biol."},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1126\/science.aea8344","article-title":"Natural grasslands used for grazing livestock can mitigate climate change","volume":"391","author":"Pillar","year":"2026","journal-title":"Science"},{"key":"B47","volume-title":"R: A Language and Environment for Statistical Computing","year":"2024"},{"key":"B48","doi-asserted-by":"publisher","first-page":"1907","DOI":"10.1890\/13-2317.1","article-title":"Higher plant diversity promotes higher diversity of fungal pathogens, while it decreases pathogen infection per plant","volume":"95","author":"Rottstock","year":"2014","journal-title":"Ecology"},{"key":"B49","doi-asserted-by":"publisher","first-page":"444","DOI":"10.2307\/4002994","article-title":"Grazing intensity effects on litter decomposition and soil nitrogen mineralization","volume":"47","author":"Shariff","year":"1994","journal-title":"J. Range Manage."},{"key":"B50","doi-asserted-by":"publisher","first-page":"1736","DOI":"10.1111\/gcb.12180","article-title":"Effects of experimental warming on fungal disease progress in oilseed rape","volume":"19","author":"Siebold","year":"2013","journal-title":"Glob. Change Biol."},{"key":"B51","doi-asserted-by":"publisher","first-page":"640","DOI":"10.1038\/s41579-023-00900-7","article-title":"Climate change impacts on plant pathogens, food security and paths forward","volume":"21","author":"Singh","year":"2023","journal-title":"Nat. Rev. Microbiol."},{"key":"B52","doi-asserted-by":"publisher","first-page":"887","DOI":"10.1016\/j.tim.2025.03.013","article-title":"Plant pathogens, microbiomes, and soil health","volume":"33","author":"Singh","year":"2025","journal-title":"Trends Microbiol."},{"key":"B53","doi-asserted-by":"publisher","first-page":"663","DOI":"10.1146\/annurev.ento.51.110104.151117","article-title":"Plant-mediated interactions between pathogenic microorganisms and herbivorous arthropods","volume":"51","author":"Stout","year":"2006","journal-title":"Annu. Rev. Entomol."},{"key":"B54","doi-asserted-by":"publisher","first-page":"1075","DOI":"10.1890\/08-2347.1","article-title":"Salicylate\u2010mediated interactions between pathogens and herbivores","volume":"91","author":"Thaler","year":"2010","journal-title":"Ecology"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1111\/1365-2664.70405","article-title":"Context-dependent responses of soil microbes and nematodes to grazing: Evidence from a global meta-analysis","volume":"63","author":"Tian","year":"2026","journal-title":"J. Appl. Ecol."},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.3390\/jof9090949","article-title":"Effects of different livestock grazing on foliar fungal diseases in an alpine grassland on the Qinghai\u2013Tibet Plateau","volume":"9","author":"Tian","year":"2023","journal-title":"J. Fungi"},{"key":"B57","doi-asserted-by":"publisher","first-page":"961","DOI":"10.1111\/ppa.12014","article-title":"Fertilization affects severity of disease caused by fungal plant pathogens","volume":"62","author":"Veresoglou","year":"2013","journal-title":"Plant Pathol."},{"key":"B58","doi-asserted-by":"publisher","first-page":"2607","DOI":"10.1094\/PDIS-07-25-1389-PDN","article-title":"First report of Nectriella adonidis causing root rot of Lespedeza chinensis in China","volume":"109","author":"Wang","year":"2025","journal-title":"Plant Dis."},{"key":"B59","doi-asserted-by":"publisher","first-page":"3181","DOI":"10.1111\/1365-2435.70169","article-title":"Arbuscular mycorrhizal fungus alters the phyllosphere microbial community and modifies plant defences against co\u2010attack of pea aphid and pathogen","volume":"39","author":"Wang","year":"2025","journal-title":"Funct. Ecol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"960","DOI":"10.1111\/gcb.14841","article-title":"Grazing simplifies soil micro\u2010food webs and decouples their relationships with ecosystem functions in grasslands","volume":"26","author":"Wang","year":"2020","journal-title":"Glob. Change Biol."},{"key":"B61","doi-asserted-by":"publisher","first-page":"1568","DOI":"10.1038\/s42003-024-07280-5","article-title":"Long-term grazing effects on soil-borne pathogens are driven by temperature","volume":"7","author":"Wang","year":"2024","journal-title":"Commun. Biol."},{"key":"B62","first-page":"315","article-title":"Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics","volume-title":"PCR Protocols: A Guide to Methods and Applications","author":"White","year":"1990"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1111\/nph.71254","article-title":"Root exudate\u2010associated microbiome assembly contributes to viral disease resistance in wheat","volume":"251","author":"Wu","year":"2026","journal-title":"New Phytol"},{"key":"B64","doi-asserted-by":"publisher","first-page":"wraf283","DOI":"10.1093\/ismejo\/wraf283","article-title":"Metabolite interactions mediate beneficial alliances between Bacillus and Trichoderma for effective Fusarium wilt control","volume":"20","author":"Xie","year":"2026","journal-title":"ISME J."},{"key":"B65","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41579-026-01290-2","article-title":"Impacts of climate extremes on plant pathogens, microbiomes and plant health","author":"Xiong","year":"2026","journal-title":"Nat. Rev. Microbiol."},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1016\/j.agee.2019.106628","article-title":"Different responses of soil element contents and their stoichiometry (C: N: P) to yak grazing and Tibetan sheep grazing in an alpine grassland on the eastern Qinghai\u2013Tibetan Plateau","volume":"285","author":"Yang","year":"2019","journal-title":"Agric. Ecosyst. Environ."},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1016\/j.agee.2022.108176","article-title":"Soil acidification caused by excessive application of nitrogen fertilizer aggravates soil-borne diseases: Evidence from literature review and field trials","volume":"340","author":"Zhang","year":"2022","journal-title":"Agric. Ecosyst. Environ."},{"key":"B68","doi-asserted-by":"publisher","first-page":"1167","DOI":"10.1111\/gcb.13431","article-title":"Grazing intensity significantly affects belowground carbon and nitrogen cycling in grassland ecosystems: A meta\u2010analysis","volume":"23","author":"Zhou","year":"2017","journal-title":"Glob. Change Biol."},{"key":"B69","doi-asserted-by":"publisher","first-page":"1301","DOI":"10.1186\/s12870-025-07356-2","article-title":"Light grazing enhances microbial-mediated nitrogen transformation in desert steppe soils","volume":"25","author":"Zhu","year":"2025","journal-title":"BMC Plant Biol."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1900475\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T05:30:31Z","timestamp":1785735031000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1900475\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,3]]},"references-count":69,"alternative-id":["10.3389\/fpls.2026.1900475"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1900475","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,3]]},"article-number":"1900475"}}