{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T21:49:41Z","timestamp":1792705781433,"version":"4.5.0"},"reference-count":73,"publisher":"Springer Science and Business Media LLC","issue":"8","license":[{"start":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T00:00:00Z","timestamp":1753142400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T00:00:00Z","timestamp":1753142400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100002347","name":"Bundesministerium f\u00fcr Bildung und Forschung","doi-asserted-by":"publisher","award":["DPPN-031A053C"],"award-info":[{"award-number":["DPPN-031A053C"]}],"id":[{"id":"10.13039\/501100002347","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004543","name":"China Scholarship Council","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004543","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat. Plants"],"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:p>Soil-borne microorganisms can systemically affect shoot resistance to pathogens relying on jasmonic acid and\/or salicylic acid. However, the emanating root triggers in these scenarios remain elusive. Here we identify an <jats:italic>N<\/jats:italic>-hydroxypipecolic-acid-(NHP-)directed, salicylic-acid-related mechanism of root-triggered systemic resistance in <jats:italic>Arabidopsis<\/jats:italic>, which uses components of systemic acquired resistance known in leaves. However, in contrast to the inductive nature of systemic acquired resistance, FLAVIN-DEPENDENT MONOOXYGENASE 1 (FMO1) continuously synthesizes NHP in roots, while the glucosyltransferase UGT76B1 concomitantly conjugates and immobilizes NHP. Physical grafting experiments and tissue-specific knockouts revealed that the loss of UGT76B1 in roots leads to enhanced NHP release, initiating shoot responses. This counteracting standby FMO1\/UGT76B1 circuit is specifically and sensitively modulated by root-associated microorganisms. Endophytic and (hemi)biotrophic fungi induce UGT76B1 degradation and FMO1 expression, resulting in varying levels of NHP being released to the shoot, where this root signal differently modulates defence and growth.<\/jats:p>","DOI":"10.1038\/s41477-025-02053-2","type":"journal-article","created":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T09:02:51Z","timestamp":1753174971000},"page":"1658-1669","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":16,"title":["A root-based N-hydroxypipecolic acid standby circuit to direct immunity and growth of Arabidopsis shoots"],"prefix":"10.1038","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-3550-5955","authenticated-orcid":false,"given":"Ping","family":"Xu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sophia","family":"Fundneider","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Birgit","family":"Lange","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rafa\u0142","family":"Maksym","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Johannes","family":"Stuttmann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9424-5548","authenticated-orcid":false,"given":"Anton R.","family":"Sch\u00e4ffner","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,7,22]]},"reference":[{"key":"2053_CR1","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1146\/annurev-phyto-082712-102340","volume":"52","author":"CMJ Pieterse","year":"2014","unstructured":"Pieterse, C. M. J. et al. Induced systemic resistance by beneficial microbes. Annu. Rev. Phytopathol. 52, 347\u2013375 (2014).","journal-title":"Annu. Rev. Phytopathol."},{"key":"2053_CR2","doi-asserted-by":"publisher","first-page":"685","DOI":"10.1016\/j.tplants.2021.01.001","volume":"26","author":"J De Kesel","year":"2021","unstructured":"De Kesel, J. et al. The induced resistance lexicon: do\u2019s and don\u2019ts. Trends Plant Sci. 26, 685\u2013691 (2021).","journal-title":"Trends Plant Sci."},{"key":"2053_CR3","doi-asserted-by":"publisher","first-page":"839","DOI":"10.1146\/annurev-arplant-042811-105606","volume":"64","author":"ZQ Fu","year":"2013","unstructured":"Fu, Z. Q. & Dong, X. Systemic acquired resistance: turning local infection into global defense. Annu. Rev. Plant Biol. 64, 839\u2013863 (2013).","journal-title":"Annu. Rev. Plant Biol."},{"key":"2053_CR4","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1146\/annurev.phyto.42.040803.140421","volume":"42","author":"WE Durrant","year":"2004","unstructured":"Durrant, W. E. & Dong, X. Systemic acquired resistance. Annu. Rev. Phytopathol. 42, 185\u2013209 (2004).","journal-title":"Annu. Rev. Phytopathol."},{"key":"2053_CR5","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1146\/annurev-phyto-080614-120132","volume":"53","author":"U Conrath","year":"2015","unstructured":"Conrath, U., Beckers, G. J. M., Langenbach, C. J. G. & Jaskiewicz, M. R. Priming for enhanced defense. Annu. Rev. Phytopathol. 53, 97\u2013119 (2015).","journal-title":"Annu. Rev. Phytopathol."},{"key":"2053_CR6","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1016\/j.pbi.2008.05.003","volume":"11","author":"AC Vlot","year":"2008","unstructured":"Vlot, A. C., Klessig, D. F. & Park, S.-W. Systemic acquired resistance: the elusive signal(s). Curr. Opin. Plant Biol. 11, 436\u2013442 (2008).","journal-title":"Curr. Opin. Plant Biol."},{"key":"2053_CR7","doi-asserted-by":"publisher","first-page":"1234","DOI":"10.1111\/nph.16953","volume":"229","author":"AC Vlot","year":"2021","unstructured":"Vlot, A. C. et al. Systemic propagation of immunity in plants. N. Phytol. 229, 1234\u20131250 (2021).","journal-title":"N. Phytol."},{"key":"2053_CR8","doi-asserted-by":"publisher","first-page":"1451","DOI":"10.1093\/plcell\/koad329","volume":"36","author":"SH Spoel","year":"2024","unstructured":"Spoel, S. H. & Dong, X. Salicylic acid in plant immunity and beyond. Plant Cell 36, 1451\u20131464 (2024).","journal-title":"Plant Cell"},{"key":"2053_CR9","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1016\/S1360-1385(02)02244-6","volume":"7","author":"U Conrath","year":"2002","unstructured":"Conrath, U., Pieterse, C. M. J. & Mauch-Mani, B. Priming in plant\u2013pathogen interactions. Trends Plant Sci. 7, 210\u2013216 (2002).","journal-title":"Trends Plant Sci."},{"key":"2053_CR10","doi-asserted-by":"publisher","first-page":"102050","DOI":"10.1016\/j.pbi.2021.102050","volume":"62","author":"J Zeier","year":"2021","unstructured":"Zeier, J. Metabolic regulation of systemic acquired resistance. Curr. Opin. Plant Biol. 62, 102050 (2021).","journal-title":"Curr. Opin. Plant Biol."},{"key":"2053_CR11","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1007\/s11104-020-04786-9","volume":"461","author":"CMJ Pieterse","year":"2021","unstructured":"Pieterse, C. M. J. et al. Pseudomonas simiae WCS417: star track of a model beneficial rhizobacterium. Plant Soil 461, 245\u2013263 (2021).","journal-title":"Plant Soil"},{"key":"2053_CR12","doi-asserted-by":"publisher","first-page":"1554","DOI":"10.4161\/psb.6.10.17443","volume":"6","author":"HA Contreras-Cornejo","year":"2011","unstructured":"Contreras-Cornejo, H. A., Mac\u00edas-Rodr\u00edguez, L., Beltr\u00e1n-Pe\u00f1a, E., Herrera-Estrella, A. & L\u00f3pez-Bucio, J. Trichoderma-induced plant immunity likely involves both hormonal- and camalexin-dependent mechanisms in Arabidopsis thaliana and confers resistance against necrotrophic fungi Botrytis cinerea. Plant Signal. Behav. 6, 1554\u20131563 (2011).","journal-title":"Plant Signal. Behav."},{"key":"2053_CR13","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1007\/s10658-011-9782-6","volume":"131","author":"MA Salas-Marina","year":"2011","unstructured":"Salas-Marina, M. A. et al. Colonization of Arabidopsis roots by Trichoderma atroviride promotes growth and enhances systemic disease resistance through jasmonic acid\/ethylene and salicylic acid pathways. Eur. J. Plant Pathol. 131, 15\u201326 (2011).","journal-title":"Eur. J. Plant Pathol."},{"key":"2053_CR14","doi-asserted-by":"publisher","first-page":"2173","DOI":"10.1104\/pp.112.207324","volume":"160","author":"JE van de Mortel","year":"2012","unstructured":"van de Mortel, J. E. et al. Metabolic and transcriptomic changes induced in Arabidopsis by the rhizobacterium Pseudomonas fluorescens SS101. Plant Physiol. 160, 2173\u20132188 (2012).","journal-title":"Plant Physiol."},{"key":"2053_CR15","doi-asserted-by":"publisher","first-page":"555","DOI":"10.1094\/MPMI-18-0555","volume":"18","author":"SE Tjamos","year":"2005","unstructured":"Tjamos, S. E., Flemetakis, E., Paplomatas, E. J. & Katinakis, P. Induction of resistance to Verticillium dahliae in Arabidopsis thaliana by the biocontrol agent K-165 and pathogenesis-related proteins gene expression. Mol. Plant Microbe Interact. 18, 555\u2013561 (2005).","journal-title":"Mol. Plant Microbe Interact."},{"key":"2053_CR16","doi-asserted-by":"publisher","first-page":"206","DOI":"10.3389\/fpls.2013.00206","volume":"4","author":"A Mart\u00ednez-Medina","year":"2013","unstructured":"Mart\u00ednez-Medina, A. et al. Deciphering the hormonal signalling network behind the systemic resistance induced by Trichoderma harzianum in tomato. Front. Plant Sci. 4, 206 (2013).","journal-title":"Front. Plant Sci."},{"key":"2053_CR17","doi-asserted-by":"publisher","first-page":"108","DOI":"10.3389\/fpls.2012.00108","volume":"3","author":"J Mathys","year":"2012","unstructured":"Mathys, J. et al. Genome-wide characterization of ISR induced in Arabidopsis thaliana by Trichoderma hamatum T382 against Botrytis cinerea infection. Front. Plant Sci. 3, 108 (2012).","journal-title":"Front. Plant Sci."},{"key":"2053_CR18","doi-asserted-by":"publisher","first-page":"e14483","DOI":"10.1111\/ppl.14483","volume":"176","author":"A Sommer","year":"2024","unstructured":"Sommer, A. et al. A salicylic acid\u2010associated plant\u2013microbe interaction attracts beneficial Flavobacterium sp. to the Arabidopsis thaliana phyllosphere. Physiol. Plant. 176, e14483 (2024).","journal-title":"Physiol. Plant."},{"key":"2053_CR19","doi-asserted-by":"publisher","first-page":"765","DOI":"10.1094\/MPMI-09-11-0253","volume":"25","author":"DJ Weston","year":"2012","unstructured":"Weston, D. J. et al. Pseudomonas fluorescens induces strain-dependent and strain-independent host plant responses in defense networks, primary metabolism, photosynthesis, and fitness. Mol. Plant Microbe Interact. 25, 765\u2013778 (2012).","journal-title":"Mol. Plant Microbe Interact."},{"key":"2053_CR20","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms11362","volume":"7","author":"S Hacquard","year":"2016","unstructured":"Hacquard, S. et al. Survival trade-offs in plant roots during colonization by closely related beneficial and pathogenic fungi. Nat. Commun. 7, 11362 (2016).","journal-title":"Nat. Commun."},{"key":"2053_CR21","doi-asserted-by":"publisher","first-page":"3367","DOI":"10.1111\/pce.14420","volume":"45","author":"M P\u00e9rez\u2010Alonso","year":"2022","unstructured":"P\u00e9rez\u2010Alonso, M. et al. The calcium sensor CBL7 is required for Serendipita indica\u2010induced growth stimulation in Arabidopsis thaliana, controlling defense against the endophyte and K+ homoeostasis in the symbiosis. Plant Cell Environ. 45, 3367\u20133382 (2022).","journal-title":"Plant Cell Environ."},{"key":"2053_CR22","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1094\/MPMI-08-22-0166-SC","volume":"36","author":"D Mart\u00ednez-Soto","year":"2023","unstructured":"Mart\u00ednez-Soto, D., Yu, H., Allen, K. S. & Ma, L.-J. Differential colonization of the plant vasculature between endophytic versus pathogenic Fusarium oxysporum strains. Mol. Plant Microbe Interact. 36, 4\u201313 (2023).","journal-title":"Mol. Plant Microbe Interact."},{"key":"2053_CR23","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1003221","volume":"9","author":"Y Brotman","year":"2013","unstructured":"Brotman, Y. et al. Trichoderma-plant root colonization: escaping early plant defense responses and activation of the antioxidant machinery for saline stress tolerance. PLoS Path. 9, e1003221 (2013).","journal-title":"PLoS Path."},{"key":"2053_CR24","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1105\/tpc.15.00496","volume":"28","author":"F Bernsdorff","year":"2016","unstructured":"Bernsdorff, F. et al. Pipecolic acid orchestrates plant systemic acquired resistance and defense priming via salicylic acid-dependent and -independent pathways. Plant Cell 28, 102\u2013129 (2016).","journal-title":"Plant Cell"},{"key":"2053_CR25","doi-asserted-by":"publisher","first-page":"500","DOI":"10.1111\/j.1365-313X.2007.03067.x","volume":"50","author":"TE Mishina","year":"2007","unstructured":"Mishina, T. E. & Zeier, J. Pathogen\u2010associated molecular pattern recognition rather than development of tissue necrosis contributes to bacterial induction of systemic acquired resistance in Arabidopsis. Plant J. 50, 500\u2013513 (2007).","journal-title":"Plant J."},{"key":"2053_CR26","doi-asserted-by":"publisher","first-page":"456","DOI":"10.1016\/j.cell.2018.02.049","volume":"173","author":"M Hartmann","year":"2018","unstructured":"Hartmann, M. et al. Flavin monooxygenase-generated N-hydroxypipecolic acid is a critical element of plant systemic immunity. Cell 173, 456\u2013469 (2018).","journal-title":"Cell"},{"key":"2053_CR27","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1038\/nature00962","volume":"419","author":"AM Maldonado","year":"2002","unstructured":"Maldonado, A. M., Doerner, P., Dixon, R. A., Lamb, C. J. & Cameron, R. K. A putative lipid transfer protein involved in systemic resistance signalling in Arabidopsis. Nature 419, 399\u2013403 (2002).","journal-title":"Nature"},{"key":"2053_CR28","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1126\/science.1170025","volume":"324","author":"HW Jung","year":"2009","unstructured":"Jung, H. W., Tschaplinski, T. J., Wang, L., Glazebrook, J. & Greenberg, J. T. Priming in systemic plant immunity. Science 324, 89\u201391 (2009).","journal-title":"Science"},{"key":"2053_CR29","doi-asserted-by":"publisher","first-page":"421","DOI":"10.1038\/ng.798","volume":"43","author":"B Chanda","year":"2011","unstructured":"Chanda, B. et al. Glycerol-3-phosphate is a critical mobile inducer of systemic immunity in plants. Nat. Genet. 43, 421\u2013427 (2011).","journal-title":"Nat. Genet."},{"key":"2053_CR30","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-11798-2","volume":"10","author":"M Wenig","year":"2019","unstructured":"Wenig, M. et al. Systemic acquired resistance networks amplify airborne defense cues. Nat. Commun. 10, 3813 (2019).","journal-title":"Nat. Commun."},{"key":"2053_CR31","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1126\/science.1147113","volume":"318","author":"S-W Park","year":"2007","unstructured":"Park, S.-W., Kaimoyo, E., Kumar, D., Mosher, S. & Klessig, D. F. Methyl salicylate is a critical mobile signal for plant systemic acquired resistance. Science 318, 113\u2013116 (2007).","journal-title":"Science"},{"key":"2053_CR32","doi-asserted-by":"publisher","first-page":"440","DOI":"10.1016\/j.molp.2020.12.018","volume":"14","author":"J Cai","year":"2021","unstructured":"Cai, J. et al. Glycosylation of N-hydroxy-pipecolic acid equilibrates between systemic acquired resistance response and plant growth. Mol. Plant 14, 440\u2013455 (2021).","journal-title":"Mol. Plant"},{"key":"2053_CR33","doi-asserted-by":"publisher","first-page":"714","DOI":"10.1093\/plcell\/koaa044","volume":"33","author":"S Bauer","year":"2021","unstructured":"Bauer, S. et al. UGT76B1, a promiscuous hub of small molecule-based immune signaling, glucosylates N-hydroxypipecolic acid, and balances plant immunity. Plant Cell 33, 714\u2013734 (2021).","journal-title":"Plant Cell"},{"key":"2053_CR34","doi-asserted-by":"publisher","first-page":"735","DOI":"10.1093\/plcell\/koaa045","volume":"33","author":"L Mohnike","year":"2021","unstructured":"Mohnike, L. et al. The glycosyltransferase UGT76B1 modulates N-hydroxy-pipecolic acid homeostasis and plant immunity. Plant Cell 33, 735\u2013749 (2021).","journal-title":"Plant Cell"},{"key":"2053_CR35","doi-asserted-by":"publisher","first-page":"750","DOI":"10.1093\/plcell\/koaa052","volume":"33","author":"EC Holmes","year":"2021","unstructured":"Holmes, E. C., Chen, Y.-C., Mudgett, M. B. & Sattely, E. S. Arabidopsis UGT76B1 glycosylates N-hydroxy-pipecolic acid and inactivates systemic acquired resistance in tomato. Plant Cell 33, 750\u2013765 (2021).","journal-title":"Plant Cell"},{"key":"2053_CR36","doi-asserted-by":"publisher","first-page":"4124","DOI":"10.1105\/tpc.111.088443","volume":"23","author":"V von Saint Paul","year":"2011","unstructured":"von Saint Paul, V. et al. The Arabidopsis glucosyltransferase UGT76B1 conjugates isoleucic acid and modulates plant defense and senescence. Plant Cell 23, 4124\u20134145 (2011).","journal-title":"Plant Cell"},{"key":"2053_CR37","doi-asserted-by":"publisher","first-page":"1679","DOI":"10.1093\/plphys\/kiab166","volume":"186","author":"I Yildiz","year":"2021","unstructured":"Yildiz, I. et al. The mobile SAR signal N-hydroxypipecolic acid induces NPR1-dependent transcriptional reprogramming and immune priming. Plant Physiol. 186, 1679\u20131705 (2021).","journal-title":"Plant Physiol."},{"key":"2053_CR38","doi-asserted-by":"publisher","first-page":"1217771","DOI":"10.3389\/fpls.2023.1217771","volume":"14","author":"M L\u00f6we","year":"2023","unstructured":"L\u00f6we, M. et al. N-hydroxypipecolic acid primes plants for enhanced microbial pattern-induced responses. Front. Plant Sci. 14, 1217771 (2023).","journal-title":"Front. Plant Sci."},{"key":"2053_CR39","doi-asserted-by":"publisher","first-page":"6444","DOI":"10.1093\/jxb\/eraa317","volume":"71","author":"A Schnake","year":"2020","unstructured":"Schnake, A. et al. Inducible biosynthesis and immune function of the systemic acquired resistance inducer N-hydroxypipecolic acid in monocotyledonous and dicotyledonous plants. J. Exp. Bot. 71, 6444\u20136459 (2020).","journal-title":"J. Exp. Bot."},{"key":"2053_CR40","doi-asserted-by":"publisher","first-page":"3795","DOI":"10.1105\/tpc.112.098343","volume":"24","author":"Y Noutoshi","year":"2012","unstructured":"Noutoshi, Y. et al. Novel plant immune-priming compounds identified via high-throughput chemical screening target salicylic acid glucosyltransferases in Arabidopsis. Plant Cell 24, 3795\u20133804 (2012).","journal-title":"Plant Cell"},{"key":"2053_CR41","doi-asserted-by":"publisher","first-page":"614","DOI":"10.1016\/j.plantsci.2005.10.016","volume":"170","author":"B Olszak","year":"2006","unstructured":"Olszak, B. et al. A putative flavin-containing mono-oxygenase as a marker for certain defense and cell death pathways. Plant Sci. 170, 614\u2013623 (2006).","journal-title":"Plant Sci."},{"key":"2053_CR42","doi-asserted-by":"publisher","first-page":"1592","DOI":"10.1093\/pcp\/pcy106","volume":"59","author":"S Joglekar","year":"2018","unstructured":"Joglekar, S. et al. Chemical activation of EDS1\/PAD4 signaling leading to pathogen resistance in Arabidopsis. Plant Cell Physiol. 59, 1592\u20131607 (2018).","journal-title":"Plant Cell Physiol."},{"key":"2053_CR43","doi-asserted-by":"publisher","first-page":"1444","DOI":"10.1104\/pp.18.01482","volume":"179","author":"KH Ryu","year":"2019","unstructured":"Ryu, K. H., Huang, L., Kang, H. M. & Schiefelbein, J. Single-cell RNA sequencing resolves molecular relationships among individual plant cells. Plant Physiol. 179, 1444\u20131456 (2019).","journal-title":"Plant Physiol."},{"key":"2053_CR44","doi-asserted-by":"publisher","first-page":"e15237","DOI":"10.1371\/journal.pone.0015237","volume":"6","author":"G Fucile","year":"2011","unstructured":"Fucile, G. et al. ePlant and the 3D data display initiative: integrative systems biology on the world wide web. PLoS ONE 6, e15237 (2011).","journal-title":"PLoS ONE"},{"key":"2053_CR45","doi-asserted-by":"publisher","first-page":"841688","DOI":"10.3389\/fpls.2022.841688","volume":"13","author":"A Shields","year":"2022","unstructured":"Shields, A., Shivnauth, V. & Castroverde, C. D. M. Salicylic acid and N-hydroxypipecolic acid at the fulcrum of the plant immunity\u2013growth equilibrium. Front. Plant Sci. 13, 841688 (2022).","journal-title":"Front. Plant Sci."},{"key":"2053_CR46","first-page":"1225","volume":"8","author":"CM Pieterse","year":"1996","unstructured":"Pieterse, C. M., Van Wees, S. C., Hoffland, E., Van Pelt, J. A. & Van Loon, L. C. Systemic resistance in Arabidopsis induced by biocontrol bacteria is independent of salicylic acid accumulation and pathogenesis-related gene expression. Plant Cell 8, 1225\u20131237 (1996).","journal-title":"Plant Cell"},{"key":"2053_CR47","doi-asserted-by":"publisher","first-page":"1571","DOI":"10.1105\/tpc.10.9.1571","volume":"10","author":"CMJ Pieterse","year":"1998","unstructured":"Pieterse, C. M. J. et al. A novel signaling pathway controlling induced systemic resistance in Arabidopsis. Plant Cell 10, 1571\u20131580 (1998).","journal-title":"Plant Cell"},{"key":"2053_CR48","doi-asserted-by":"publisher","first-page":"464","DOI":"10.1016\/j.cell.2016.02.028","volume":"165","author":"K Hiruma","year":"2016","unstructured":"Hiruma, K. et al. Root endophyte Colletotrichum tofieldiae confers plant fitness benefits that are phosphate status dependent. Cell 165, 464\u2013474 (2016).","journal-title":"Cell"},{"key":"2053_CR49","doi-asserted-by":"publisher","first-page":"e0121902","DOI":"10.1371\/journal.pone.0121902","volume":"10","author":"R Lyons","year":"2015","unstructured":"Lyons, R. et al. Fusarium oxysporum triggers tissue-specific transcriptional reprogramming in Arabidopsis thaliana. PLoS ONE 10, e0121902 (2015).","journal-title":"PLoS ONE"},{"key":"2053_CR50","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1071\/AP06060","volume":"35","author":"CI Edgar","year":"2006","unstructured":"Edgar, C. I. et al. Salicylic acid mediates resistance to the vascular wilt pathogen Fusarium oxysporum in the model host Arabidopsis thaliana. Australas. Plant Pathol. 35, 581\u2013591 (2006).","journal-title":"Australas. Plant Pathol."},{"key":"2053_CR51","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1111\/mpp.12141","volume":"15","author":"A Alonso\u2010Ram\u00edrez","year":"2014","unstructured":"Alonso\u2010Ram\u00edrez, A. et al. Salicylic acid prevents Trichoderma harzianum from entering the vascular system of roots. Mol. Plant Pathol. 15, 823\u2013831 (2014).","journal-title":"Mol. Plant Pathol."},{"key":"2053_CR52","doi-asserted-by":"publisher","first-page":"e69352","DOI":"10.1371\/journal.pone.0069352","volume":"8","author":"L Pedrotti","year":"2013","unstructured":"Pedrotti, L., Mueller, M. J. & Waller, F. Piriformospora indica root colonization triggers local and systemic root responses and inhibits secondary colonization of distal roots. PLoS ONE 8, e69352 (2013).","journal-title":"PLoS ONE"},{"key":"2053_CR53","doi-asserted-by":"publisher","first-page":"6052","DOI":"10.1093\/jxb\/erac263","volume":"73","author":"L Wang","year":"2022","unstructured":"Wang, L., Calabria, J., Chen, H.-W. & Somssich, M. The Arabidopsis thaliana\u2013Fusarium oxysporum strain 5176 pathosystem: an overview. J. Exp. Bot. 73, 6052\u20136067 (2022).","journal-title":"J. Exp. Bot."},{"key":"2053_CR54","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1146\/annurev-cellbio-092910-154055","volume":"28","author":"CMJ Pieterse","year":"2012","unstructured":"Pieterse, C. M. J., Van der Does, D., Zamioudis, C., Leon-Reyes, A. & Van Wees, S. C. M. Hormonal modulation of plant immunity. Annu. Rev. Cell Dev. Biol. 28, 489\u2013521 (2012).","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"2053_CR55","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1111\/tpj.15135","volume":"105","author":"R Eichmann","year":"2021","unstructured":"Eichmann, R., Richards, L. & Sch\u00e4fer, P. Hormones as go\u2010betweens in plant microbiome assembly. Plant J. 105, 518\u2013541 (2021).","journal-title":"Plant J."},{"key":"2053_CR56","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1038\/s41579-020-0412-1","volume":"18","author":"P Trivedi","year":"2020","unstructured":"Trivedi, P., Leach, J. E., Tringe, S. G., Sa, T. & Singh, B. K. Plant\u2013microbiome interactions: from community assembly to plant health. Nat. Rev. Microbiol. 18, 607\u2013621 (2020).","journal-title":"Nat. Rev. Microbiol."},{"key":"2053_CR57","doi-asserted-by":"publisher","first-page":"1386","DOI":"10.1890\/06-0219","volume":"88","author":"J Schimel","year":"2007","unstructured":"Schimel, J., Balser, T. C. & Wallenstein, M. Microbial stress\u2010response physiology and its implications for ecosystem function. Ecology 88, 1386\u20131394 (2007).","journal-title":"Ecology"},{"key":"2053_CR58","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1093\/jxb\/eraa417","volume":"72","author":"V Chaudhry","year":"2021","unstructured":"Chaudhry, V. et al. Shaping the leaf microbiota: plant\u2013microbe\u2013microbe interactions. J. Exp. Bot. 72, 36\u201356 (2021).","journal-title":"J. Exp. Bot."},{"key":"2053_CR59","doi-asserted-by":"publisher","first-page":"1692","DOI":"10.3389\/fpls.2018.01692","volume":"9","author":"C Chuberre","year":"2018","unstructured":"Chuberre, C. et al. Plant immunity is compartmentalized and specialized in roots. Front. Plant Sci. 9, 1692 (2018).","journal-title":"Front. Plant Sci."},{"key":"2053_CR60","doi-asserted-by":"publisher","first-page":"1666","DOI":"10.1104\/pp.106.081257","volume":"141","author":"TE Mishina","year":"2006","unstructured":"Mishina, T. E. & Zeier, J. The Arabidopsis flavin-dependent monooxygenase FMO1 is an essential component of biologically induced systemic acquired resistance. Plant Physiol. 141, 1666\u20131675 (2006).","journal-title":"Plant Physiol."},{"key":"2053_CR61","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1007\/s00299-024-03228-5","volume":"43","author":"W Zhang","year":"2024","unstructured":"Zhang, W., Maksym, R., Georgii, E., Geist, B. & Sch\u00e4ffner, A. R. SA and NHP glucosyltransferase UGT76B1 affects plant defense in both SID2- and NPR1-dependent and independent manner. Plant Cell Rep. 43, 149 (2024).","journal-title":"Plant Cell Rep."},{"key":"2053_CR62","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1016\/S0014-5793(04)00148-6","volume":"561","author":"S Bensmihen","year":"2004","unstructured":"Bensmihen, S. et al. Analysis of an activated ABI5 allele using a new selection method for transgenic Arabidopsis seeds. FEBS Lett. 561, 127\u2013131 (2004).","journal-title":"FEBS Lett."},{"key":"2053_CR63","doi-asserted-by":"publisher","first-page":"735","DOI":"10.1046\/j.1365-313x.1998.00343.x","volume":"16","author":"SJ Clough","year":"1998","unstructured":"Clough, S. J. & Bent, A. F. Floral dip: a simplified method for Agrobacterium\u2010mediated transformation of Arabidopsis thaliana. Plant J. 16, 735\u2013743 (1998).","journal-title":"Plant J."},{"key":"2053_CR64","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1111\/tpj.15197","volume":"106","author":"J Stuttmann","year":"2021","unstructured":"Stuttmann, J. et al. Highly efficient multiplex editing: one\u2010shot generation of 8\u00d7 Nicotiana benthamiana and 12\u00d7 Arabidopsis mutants. Plant J. 106, 8\u201322 (2021).","journal-title":"Plant J."},{"key":"2053_CR65","doi-asserted-by":"publisher","first-page":"839","DOI":"10.1021\/sb4001504","volume":"3","author":"C Engler","year":"2014","unstructured":"Engler, C. et al. A Golden Gate modular cloning toolbox for plants. ACS Synth. Biol. 3, 839\u2013843 (2014).","journal-title":"ACS Synth. Biol."},{"key":"2053_CR66","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1111\/tpj.13319","volume":"89","author":"J Ordon","year":"2017","unstructured":"Ordon, J. et al. Generation of chromosomal deletions in dicotyledonous plants employing a user\u2010friendly genome editing toolkit. Plant J. 89, 155\u2013168 (2017).","journal-title":"Plant J."},{"key":"2053_CR67","doi-asserted-by":"publisher","first-page":"519","DOI":"10.1111\/j.1365-313X.2009.04060.x","volume":"61","author":"TL Shimada","year":"2010","unstructured":"Shimada, T. L., Shimada, T. & Hara\u2010Nishimura, I. A rapid and non\u2010destructive screenable marker, FAST, for identifying transformed seeds of Arabidopsis thaliana. Plant J. 61, 519\u2013528 (2010).","journal-title":"Plant J."},{"key":"2053_CR68","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1046\/j.1365-313X.1996.09020195.x","volume":"9","author":"D Lagarde","year":"1996","unstructured":"Lagarde, D. et al. Tissue\u2010specific expression of Arabidopsis AKT1 gene is consistent with a role in K+ nutrition. Plant J. 9, 195\u2013203 (1996).","journal-title":"Plant J."},{"key":"2053_CR69","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1111\/j.1365-313X.2007.03234.x","volume":"52","author":"A Christmann","year":"2007","unstructured":"Christmann, A., Weiler, E. W., Steudle, E. & Grill, E. A hydraulic signal in root\u2010to\u2010shoot signalling of water shortage. Plant J. 52, 167\u2013174 (2007).","journal-title":"Plant J."},{"key":"2053_CR70","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1186\/s13007-020-00584-7","volume":"16","author":"M Osman","year":"2020","unstructured":"Osman, M., Stigloher, C., Mueller, M. J. & Waller, F. An improved growth medium for enhanced inoculum production of the plant growth-promoting fungus Serendipita indica. Plant Meth. 16, 39 (2020).","journal-title":"Plant Meth."},{"key":"2053_CR71","doi-asserted-by":"publisher","first-page":"e0039","DOI":"10.1199\/tab.0039","volume":"1","author":"F Katagiri","year":"2002","unstructured":"Katagiri, F., Thilmony, R. & He, S. Y. The Arabidopsis thaliana\u2013Pseudomonas syringae interaction. Arabidopsis Book 1, e0039 (2002).","journal-title":"Arabidopsis Book"},{"key":"2053_CR72","doi-asserted-by":"publisher","unstructured":"Sch\u00e4ffner, A. R. & Xu, P. Original LC\u2013MS and confocal data of the study \u2018A root-based N-hydroxypipecolic acid standby circuit to direct immunity and growth of Arabidopsis shoots\u2019 part 1. OSF https:\/\/doi.org\/10.17605\/OSF.IO\/HKX75 (2025).","DOI":"10.17605\/OSF.IO\/HKX75"},{"key":"2053_CR73","doi-asserted-by":"publisher","unstructured":"Sch\u00e4ffner, A. R. & Xu, P. Original LC\u2013MS and confocal data of the study \u2018A root-based N-hydroxypipecolic acid standby circuit to direct immunity and growth of Arabidopsis shoots\u2019 part 2. OSF https:\/\/doi.org\/10.17605\/OSF.IO\/EV796 (2025).","DOI":"10.17605\/OSF.IO\/EV796"}],"container-title":["Nature Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41477-025-02053-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41477-025-02053-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41477-025-02053-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,7]],"date-time":"2025-09-07T12:42:59Z","timestamp":1757248979000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41477-025-02053-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,22]]},"references-count":73,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2025,8]]}},"alternative-id":["2053"],"URL":"https:\/\/doi.org\/10.1038\/s41477-025-02053-2","relation":{},"ISSN":["2055-0278"],"issn-type":[{"value":"2055-0278","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,22]]},"assertion":[{"value":"15 November 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"13 June 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 July 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}