{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T06:12:32Z","timestamp":1788502352159,"version":"build-2803163510"},"reference-count":43,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T00:00:00Z","timestamp":1788480000000},"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:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>\n                      <jats:italic>Phellodendron chinense<\/jats:italic>\n                      is an important medicinal plant in China. However, continuous monoculture has caused soil degradation and physiological constraints, which have become a key bottleneck limiting its sustainable production. Intercropping with companion plants is an effective approach to break this dilemma.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      We selected five representative cultivation systems commonly practiced of\n                      <jats:italic>Phellodendron chinense<\/jats:italic>\n                      and systematically measured their rhizosphere soil properties, microbial community composition, and leaf gas-exchange traits. The overarching goal was twofold: (i) to characterize the intercropping-induced variations in these three dimensions (soil properties, microbial communities, and photosynthetic physiology, and (ii) to elucidate the coupling relationships among these three dimensions.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results and Discussion<\/jats:title>\n                    <jats:p>\n                      The results demonstrated that total nitrogen(N), total sodium(Na), and magnesium(Mg) contents in the T4 and T2 patterns were significantly higher than those in other patterns. The T2 and T4 patterns significantly enhanced microbial community \u03b1-diversity. In contrast, the T3 pattern caused significant soil acidification (pH 4.69) and elevated the relative abundance of the tolerant phylum Chloroflexi. T4 and T2 patterns significantly alleviated stomatal limitations in\n                      <jats:italic>P. chinense<\/jats:italic>\n                      , achieving optimal net photosynthetic rate(Pn) and stomatal conductance(Gs). Correlation analysis showed that Pn was significantly and positively correlated with soil P and Mg contents, while Gs was significantly and positively correlated with soil Na content, and N and Na were the critical factors affecting the structural variation of bacterial and fungal communities. Overall, this study demonstrates that intercropping promote soil nutrient accumulation enhances rhizosphere microbial diversity to facilitate the alleviation of photosynthetic physiological limitations. Intercropping\n                      <jats:italic>P. chinense<\/jats:italic>\n                      with bamboo or tea synergistically mitigates the host\u2019s photosynthetic physiological limitations through a belowground positive feedback pathway. These findings provide an important theoretical basis for the efficient and sustainable cultivation of this rare medicinal plant.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fpls.2026.1882609","type":"journal-article","created":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T05:29:12Z","timestamp":1788499752000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Early-stage intercropping modulates rhizosphere soil properties, microbiome composition, and photosynthetic physiology of Phellodendron chinense: a comparative analysis of five cultivation systems"],"prefix":"10.3389","volume":"17","author":[{"given":"Chuan","family":"Xie","sequence":"first","affiliation":[{"name":"Ecological Conservation, Restoration and Resource Utilization in Forest and Wetland Key Laboratory of Sichuan Province, Sichuan Academy of Forestry","place":["Chengdu, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiaojiao","family":"Wu","sequence":"additional","affiliation":[{"name":"School of Materials and Environmental Engineering, Chengdu Technological University","place":["Chengdu, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peng","family":"Song","sequence":"additional","affiliation":[{"name":"Ecological Conservation, Restoration and Resource Utilization in Forest and Wetland Key Laboratory of Sichuan Province, Sichuan Academy of Forestry","place":["Chengdu, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiuchan","family":"Xiao","sequence":"additional","affiliation":[{"name":"School of Materials and Environmental Engineering, Chengdu Technological University","place":["Chengdu, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinyuan","family":"An","sequence":"additional","affiliation":[{"name":"School of Materials and Environmental Engineering, Chengdu Technological University","place":["Chengdu, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huawei","family":"Wu","sequence":"additional","affiliation":[{"name":"Ecological Conservation, Restoration and Resource Utilization in Forest and Wetland Key Laboratory of Sichuan Province, Sichuan Academy of Forestry","place":["Chengdu, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhipeng","family":"Sun","sequence":"additional","affiliation":[{"name":"Ecological Conservation, Restoration and Resource Utilization in Forest and Wetland Key Laboratory of Sichuan Province, Sichuan Academy of Forestry","place":["Chengdu, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,4]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"444","DOI":"10.1111\/brv.12119","article-title":"Influences of evergreen gymnosperm and deciduous angiosperm tree species on the functioning of temperate and boreal forests","volume":"90","author":"Augusto","year":"2015","journal-title":"Biol. Rev."},{"key":"B2","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1093\/femsre\/fuw040","article-title":"Forest microbiome: diversity, complexity and dynamics","volume":"41","author":"Baldrian","year":"2017","journal-title":"FEMS Microbiol. Rev."},{"key":"B3","doi-asserted-by":"publisher","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":"B4","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/j.1574-6941.2009.00654.x","article-title":"Plant species and soil type cooperatively shape the structure and function of microbial communities in the rhizosphere","volume":"68","author":"Berg","year":"2009","journal-title":"FEMS Microbiol. Ecol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1104\/pp.17.01624","article-title":"The impacts of phosphorus deficiency on the photosynthetic electron transport chain","volume":"177","author":"Carstensen","year":"2018","journal-title":"Plant Physiol."},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1186\/s13750-022-00260-4","article-title":"Evidence for the impacts of agroforestry on ecosystem services and human well-being in high-income countries: a systematic map","volume":"11","author":"Castle","year":"2022","journal-title":"Environ. Evidence"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2023.1193990","article-title":"Soil chemistry, metabarcoding, and metabolome analyses reveal that a sugarcane-Dictyophora indusiata intercropping system can enhance soil health by reducing soil nitrogen loss","volume":"14","author":"Duan","year":"2023","journal-title":"Front. Microbiol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"1354","DOI":"10.1890\/05-1839","article-title":"Toward an ecological classification of soil bacteria","volume":"88","author":"Fierer","year":"2007","journal-title":"Ecology"},{"key":"B9","doi-asserted-by":"publisher","first-page":"1078","DOI":"10.1104\/pp.111.175331","article-title":"P for two, sharing a scarce resource: soil phosphorus acquisition in the rhizosphere of intercropped species","volume":"156","author":"Hinsinger","year":"2011","journal-title":"Plant Physiol."},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1186\/2049-2618-1-22","article-title":"Community genomic analyses constrain the distribution of metabolic traits across the Chloroflexi phylum and indicate roles in sediment carbon cycling","volume":"1","author":"Hug","year":"2013","journal-title":"Microbiome"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/j.indcrop.2026.122837","article-title":"Functional reshaping of rhizosphere microorganisms drives the degradation of soil multifunctionality in continuously cultivated Rhodiola crenulata","volume":"241","author":"Jia","year":"2026","journal-title":"Ind. Crops Prod."},{"key":"B12","doi-asserted-by":"publisher","first-page":"4966","DOI":"10.1038\/s41598-020-61418-z","article-title":"Rooting behaviour and soil properties in different bamboo species of Western Himalayan Foothills","volume":"10","author":"Kaushal","year":"2020","journal-title":"Sci. Rep."},{"key":"B13","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1023\/a:1026206511084","article-title":"Tannins in nutrient dynamics of forest ecosystems-a review","volume":"256","author":"Kraus","year":"2003","journal-title":"Plant Soil"},{"key":"B14","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s11104-013-1801-2","article-title":"Sodium as nutrient and toxicant","volume":"369","author":"Kronzucker","year":"2013","journal-title":"Plant Soil"},{"key":"B15","doi-asserted-by":"publisher","first-page":"3386","DOI":"10.1111\/j.1365-2486.2010.02179.x","article-title":"Time lag between photosynthesis and carbon dioxide efflux from soil: a review of mechanisms and controls","volume":"16","author":"Kuzyakov","year":"2010","journal-title":"Global Change Biol."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1016\/j.micres.2023.127375","article-title":"Roles of phosphate-solubilizing bacteria in mediating soil legacy phosphorus availability","volume":"272","author":"Li","year":"2023","journal-title":"Microbiol. Res."},{"key":"B17","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1007\/s00442-005-0256-4","article-title":"Root distribution and interactions between intercropped species","volume":"2","author":"Li","year":"2006","journal-title":"Oecologia"},{"key":"B18","doi-asserted-by":"publisher","first-page":"4365","DOI":"10.2147\/dddt.s385301","article-title":"Anti-inflammatory activation of Phellodendri Chinensis Cortex is mediated by berberine erythrocytes self-assembly targeted delivery system","volume":"16","author":"Li","year":"2022","journal-title":"Drug Des. Dev. Ther."},{"key":"B19","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1002\/jobm.200900338","article-title":"Lignin-modifying enzymes in filamentous basidiomycetes-ecological, functional and phylogenetic review","volume":"50","author":"Lundell","year":"2010","journal-title":"J. Basic Microbiol."},{"key":"B20","article-title":"Limonoids with antimicrobial activity from phellodendron chinenses chneid","volume-title":"Chemistry & Biodiversity","author":"Luo","year":"2025"},{"key":"B21","doi-asserted-by":"publisher","first-page":"1217","DOI":"10.17660\/ActaHortic.2018.1217.2","article-title":"Nutrient availability in the rhizosphere: a review","author":"Mimmo","year":"2018","journal-title":"Acta Hortic."},{"key":"B22","doi-asserted-by":"publisher","first-page":"796","DOI":"10.1080\/00380768.2011.629176","article-title":"Tea plant (Camellia sinensis L.) roots secrete oxalic acid and caffeine into medium containing aluminum","volume":"57","author":"Morita","year":"2011","journal-title":"Soil Sci. Plant Nutr."},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.3390\/ijms24054640","article-title":"Aluminum and fluoride stresses altered organic acid and secondary metabolism in tea (camellia sinensis) plants: influences on plant tolerance, tea quality and safety","volume":"24","author":"Peng","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"B24","doi-asserted-by":"publisher","first-page":"789","DOI":"10.1038\/nrmicro3109","article-title":"Going back to the roots: the microbial ecology of the rhizosphere","volume":"11","author":"Philippot,","year":"2015","journal-title":"Nature Reviews Microbiology"},{"key":"B25","doi-asserted-by":"publisher","first-page":"1340","DOI":"10.1038\/ismej.2010.58","article-title":"Soil bacterial and fungal communities across a pH gradient in an arable soil","volume":"4","author":"Rousk","year":"2010","journal-title":"ISME J."},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1016\/j.soilbio.2009.10.014","article-title":"Phenol oxidase, peroxidase and organic matter dynamics of soil[J]","volume":"2010","author":"Sinsabaugh","year":"2010","journal-title":"Soil Biology and Biochemistry"},{"key":"B27","first-page":"397","article-title":"Anti-inflammatory response and detection of pharmacologically active components in serum of rats with carrageenan-induced paw swelling treated with phellodendron chinense Revista brasileira de farmacognosia","author":"Shi","year":"2023"},{"key":"B28","doi-asserted-by":"publisher","first-page":"992","DOI":"10.1038\/ismej.2009.43","article-title":"Abundance, composition, diversity and novelty of soil Proteobacteria","volume":"3","author":"Spain","year":"2009","journal-title":"ISME J."},{"key":"B29","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1016\/j.scitotenv.2018.07.038","article-title":"Leachate from fine root litter is more acidic than leaf litter leachate: a 2.5-year laboratory incubation","volume":"645","author":"Tanikawa","year":"2018","journal-title":"Sci. Total Environ."},{"key":"B30","first-page":"1256688","article-title":"Global diversity and geography of soil fungi","volume-title":"Science","author":"Tedersoo","year":"2014"},{"key":"B31","first-page":"414","article-title":"Functioning of potassium and magnesium in photosynthesis, photosynthate translocation and photoprotection","volume-title":"Physiologia Plantarum.","author":"Tr\u00e4nkner","year":"2018"},{"key":"B32","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1007\/bf00346060","article-title":"Nutrient immobilization and release patterns during plant decomposition in a dry tropical bamboo savanna, India","volume":"14","author":"Tripathi","year":"1992","journal-title":"Biol. Fertil. Soils"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.3389\/fenvs.2019.00168","article-title":"The role of plant litter in driving plant-soil feedbacks","volume":"7","author":"Veen","year":"2019","journal-title":"Front. Environ. Sci."},{"key":"B34","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1111\/j.1469-8137.2012.04190.x","article-title":"Opportunities for improving phosphorus-use efficiency in crop plants","volume":"195","author":"Veneklaas","year":"2012","journal-title":"New Phytol."},{"key":"B35","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1038\/ismej.2012.116","article-title":"Fungal community on decomposing leaf litter undergoes rapid successional changes","volume":"7","author":"Vo\u0159\u00ed\u0161kov\u00e1","year":"2013","journal-title":"ISME J."},{"key":"B36","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-540-95966-3","volume-title":"Tree and Forest Measurement","author":"West","year":"2009"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1016\/j.soilbio.2020.107953","article-title":"Soil microbial diversity and composition: links to soil texture and associated properties","volume":"149","author":"Xia","year":"2020","journal-title":"Soil Biol. Biochem."},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecoenv.2025.117703","article-title":"Exploring the link between soil health and crop productivity","volume":"289","author":"Xing","year":"2025","journal-title":"Ecotoxicology Environ. Saf."},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2022.735359","article-title":"Fine-root decomposition and nutrient return in moso bamboo (phyllostachys pubescens J.Houz.) plantations in southeast China","volume":"13","author":"Xu","year":"2022","journal-title":"Front. Plant Sci."},{"key":"B40","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1016\/j.geoderma.2019.02.017","article-title":"Responses of soil organic carbon decomposition to warming depend on the natural warming gradient","volume":"343","author":"Xu","year":"2019","journal-title":"Geoderma"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1016\/j.soilbio.2021.108399","article-title":"Dominant plants affect litter decomposition mainly through modifications of the soil microbial community","volume":"161","author":"Yang","year":"2021","journal-title":"Soil Biol. Biochem."},{"key":"B42","doi-asserted-by":"publisher","first-page":"1663","DOI":"10.1007\/s11104-025-07526-z","article-title":"Meta-analysis of soil oxygen distribution: plant species-specific dynamics and optimal levels for plant growth","volume":"514","author":"Zhang","year":"2025","journal-title":"Plant Soil"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.1083788","article-title":"Berberine a traditional Chinese drug repurposing: its actions in inflammation-associated ulcerative colitis and cancer therapy","volume":"13","author":"Zhu","year":"2022","journal-title":"Front. Immunol."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1882609\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T05:29:13Z","timestamp":1788499753000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1882609\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,4]]},"references-count":43,"alternative-id":["10.3389\/fpls.2026.1882609"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1882609","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,4]]},"article-number":"1882609"}}