{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,31]],"date-time":"2026-08-31T08:19:50Z","timestamp":1788164390521,"version":"build-2803163510"},"reference-count":73,"publisher":"American Chemical Society (ACS)","issue":"8","license":[{"start":{"date-parts":[[2026,3,16]],"date-time":"2026-03-16T00:00:00Z","timestamp":1773619200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP19H05681"],"award-info":[{"award-number":["JP19H05681"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP25K09669"],"award-info":[{"award-number":["JP25K09669"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP25K09683"],"award-info":[{"award-number":["JP25K09683"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,8,24]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Arbuscular mycorrhizal (AM) fungi form mutualistic symbioses with a wide range of terrestrial plants, in which they exchange inorganic nutrients taken up from the soil for plant-derived photosynthates such as sugars and lipids. This nutrient exchange occurs exclusively at arbuscules\u2500highly branched hyphal structures that develop in the plant root cortex. Although the molecular mechanisms underlying arbuscule development and function have been extensively studied using targeted approaches based on transgenic plants and fluorescence microscopy, directly visualizing the various molecular components crucial for arbuscule function using nontargeted approaches remains a major challenge. Here, we demonstrate that ultrabroadband multiplex coherent anti-Stokes Raman scattering (CARS) microspectroscopy enables two- and three-dimensional imaging of arbuscule-containing regions in plant roots. Using genetically transformed Lotus japonicus roots colonized by the AM fungus Rhizophagus irregularis, we identify Raman spectroscopic signatures of arbuscules indicative of protein-rich domains. These domains are associated with periarbuscular membrane-embedded transporters that are involved in AM nutrient exchange, including phosphate transporters, as well as other proteins present in fungal and plant cells adjacent to the arbuscules. Additionally, we detect adjacent lipid-rich regions corresponding to arbuscule trunks that mainly contain unsaturated triacylglycerols transferred from the host plant. Our findings highlight the potential of ultrabroadband multiplex CARS imaging as a label-free, in situ imaging tool for studying arbuscules, providing deeper chemical insights into AM symbiosis.<\/jats:p>","DOI":"10.1021\/cbmi.6c00006","type":"journal-article","created":{"date-parts":[[2026,3,16]],"date-time":"2026-03-16T21:54:21Z","timestamp":1773698061000},"page":"1794-1804","source":"Crossref","is-referenced-by-count":0,"title":["Ultrabroadband Multiplex\nCARS Imaging Reveals Distinct\nProtein- and Lipid-Rich Domains in Arbuscules"],"prefix":"10.1021","volume":"4","author":[{"given":"Amu","family":"Sofue","sequence":"first","affiliation":[{"name":"Kwansei Gakuin University , , 1 Gakuen Uegahara , ,","place":["Sanda, Japan, 669-1330"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Naoya","family":"Takeda","sequence":"additional","affiliation":[{"name":"Kwansei Gakuin University , , 1 Gakuen Uegahara , ,","place":["Sanda, Japan, 669-1330"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2035-2068","authenticated-orcid":true,"given":"Shinsuke","family":"Shigeto","sequence":"additional","affiliation":[{"name":"Kwansei Gakuin University , , 1 Gakuen Uegahara , ,","place":["Sanda, Japan, 669-1330"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,3,16]]},"reference":[{"key":"2026083104104745000_cit1","volume-title":"Mycorrhizal Symbiosis","author":"Smith","year":"2008"},{"key":"2026083104104745000_cit2","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1038\/ncomms1046","article-title":"Mechanisms Underlying\nBeneficial Plant\u2013Fungus Interactions\nin Mycorrhizal Symbiosis","volume":"1","author":"Bonfante","year":"2010","journal-title":"Nat. Commun."},{"key":"2026083104104745000_cit3","doi-asserted-by":"publisher","first-page":"1108","DOI":"10.1111\/nph.14976","article-title":"Evolutionary History of Mycorrhizal Symbioses and Global\nHost Plant Diversity","volume":"220","author":"Brundrett","year":"2018","journal-title":"New Phytol."},{"key":"2026083104104745000_cit4","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1146\/annurev-arplant-042110-103846","article-title":"Roles of Arbuscular Mycorrhizas in Plant Nutrition\nand Growth: New Paradigms from Cellular to Ecosystem Scales","volume":"62","author":"Smith","year":"2011","journal-title":"Annu. Rev. Plant Biol."},{"key":"2026083104104745000_cit5","doi-asserted-by":"publisher","DOI":"10.1126\/science.aba1223","article-title":"How Mycorrhizal\nAssociations Drive\nPlant Population and Community Biology","volume":"367","author":"Tedersoo","year":"2020","journal-title":"Science"},{"key":"2026083104104745000_cit6","doi-asserted-by":"publisher","first-page":"R952","DOI":"10.1016\/j.cub.2017.06.042","article-title":"Understanding the Arbuscule at the\nHeart of Endomycorrhizal\nSymbioses in Plants","volume":"27","author":"Luginbuehl","year":"2017","journal-title":"Curr. Biol."},{"key":"2026083104104745000_cit7","doi-asserted-by":"publisher","first-page":"194","DOI":"10.1038\/s41477-019-0364-5","article-title":"Extensive Membrane\nSystems at the Host\u2013Arbuscular Mycorrhizal Fungus Interface","volume":"5","author":"Ivanov","year":"2019","journal-title":"Nat. Plants"},{"key":"2026083104104745000_cit8","doi-asserted-by":"publisher","first-page":"809","DOI":"10.1104\/pp.109.141879","article-title":"Live-Cell Imaging\nReveals Periarbuscular Membrane Domains\nand Organelle Location in Medicago truncatula Roots\nDuring Arbuscular Mycorrhizal Symbiosis","volume":"151","author":"Pumplin","year":"2009","journal-title":"Plant\nPhysiol."},{"key":"2026083104104745000_cit9","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1146\/annurev-cellbio-101512-122413","article-title":"Cell and Developmental\nBiology of Arbuscular Mycorrhiza\nSymbiosis","volume":"29","author":"Gutjahr","year":"2013","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"2026083104104745000_cit10","doi-asserted-by":"publisher","first-page":"2319","DOI":"10.1105\/tpc.17.00555","article-title":"Plant Signaling\nand Metabolic Pathways\nEnabling Arbuscular Mycorrhizal Symbiosis","volume":"29","author":"MacLean","year":"2017","journal-title":"Plant\nCell"},{"key":"2026083104104745000_cit11","doi-asserted-by":"publisher","first-page":"1151","DOI":"10.1111\/tpj.12706","article-title":"A Set of Fluorescent\nProtein-Based Markers Expressed\nfrom Constitutive and Arbuscular Mycorrhiza-Inducible Promoters to\nLabel Organelles, Membranes and Cytoskeletal Elements in Medicago truncatula","volume":"80","author":"Ivanov","year":"2014","journal-title":"Plant J."},{"key":"2026083104104745000_cit12","doi-asserted-by":"publisher","first-page":"19651","DOI":"10.1021\/jacs.2c05359","article-title":"Raman Spectroscopy for Chemical Biology\nResearch","volume":"144","author":"Dodo","year":"2022","journal-title":"J. Am. Chem. Soc."},{"key":"2026083104104745000_cit13","doi-asserted-by":"publisher","first-page":"5661","DOI":"10.1021\/ac300834f","article-title":"Disentangling Dynamic\nChanges of Multiple Cellular Components During the Yeast Cell Cycle\nby in Vivo Multivariate Raman Imaging","volume":"84","author":"Huang","year":"2012","journal-title":"Anal. Chem."},{"key":"2026083104104745000_cit14","doi-asserted-by":"publisher","first-page":"12501","DOI":"10.1021\/acs.analchem.9b03261","article-title":"Inhomogeneous Molecular Distributions\nand Cytochrome Types and Redox States in Fungal Cells Revealed by\nRaman Hyperspectral Imaging Using Multivariate Curve Resolution\u2013Alternating\nLeast Squares","volume":"91","author":"Yasuda","year":"2019","journal-title":"Anal. Chem."},{"key":"2026083104104745000_cit15","doi-asserted-by":"publisher","first-page":"ME22006","DOI":"10.1264\/jsme2.me22006","article-title":"Raman Microspectroscopy and Imaging\nof Filamentous\nFungi","volume":"37","author":"Shigeto","year":"2022","journal-title":"Microbes Environ."},{"key":"2026083104104745000_cit16","doi-asserted-by":"publisher","first-page":"7295","DOI":"10.1021\/ac401188f","article-title":"Correlation of Carotenoid\nAccumulation with Aggregation and Biofilm Development in Rhodococcus sp. SD-74","volume":"85","author":"Zheng","year":"2013","journal-title":"Anal. Chem."},{"key":"2026083104104745000_cit17","doi-asserted-by":"publisher","first-page":"7704","DOI":"10.1038\/s41598-020-64737-3","article-title":"Raman Spectroscopic\nSignatures of Carotenoids and Polyenes Enable Label-Free Visualization\nof Microbial Distributions within Pink Biofilms","volume":"10","author":"Horiue","year":"2020","journal-title":"Sci. Rep."},{"key":"2026083104104745000_cit18","doi-asserted-by":"publisher","first-page":"5055","DOI":"10.1021\/ac400266a","article-title":"Label-Free\nChemically Specific Imaging in Planta with Stimulated\nRaman Scattering Microscopy","volume":"85","author":"Mansfield","year":"2013","journal-title":"Anal. Chem."},{"key":"2026083104104745000_cit19","doi-asserted-by":"publisher","first-page":"9436","DOI":"10.1021\/acs.analchem.5b02434","article-title":"Vibrational Fingerprint Mapping Reveals\nSpatial Distribution of Functional Groups of Lignin in Plant Cell\nWall","volume":"87","author":"Liu","year":"2015","journal-title":"Anal. Chem."},{"key":"2026083104104745000_cit20","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1016\/j.jplph.2017.11.002","article-title":"In Vivo Cytological and Chemical Analysis of Casparian Strips\nUsing Stimulated Raman Scattering Microscopy","volume":"220","author":"Man","year":"2018","journal-title":"J. Plant Physiol."},{"key":"2026083104104745000_cit21","doi-asserted-by":"publisher","first-page":"490","DOI":"10.3389\/fpls.2014.00490","article-title":"1064 nm\nFT-Raman Spectroscopy for Investigations of Plant Cell Walls and Other\nBiomass Materials","volume":"5","author":"Agarwal","year":"2014","journal-title":"Front. Plant Sci."},{"key":"2026083104104745000_cit22","doi-asserted-by":"publisher","first-page":"10","DOI":"10.3389\/fchem.2016.00010","article-title":"Waterproofing\nin Arabidopsis: Following Phenolics and Lipids in Situ by Confocal Raman Microscopy","volume":"4","author":"Prats Mateu","year":"2016","journal-title":"Front. Chem."},{"key":"2026083104104745000_cit23","doi-asserted-by":"publisher","first-page":"1234","DOI":"10.1039\/D0AN02181D","article-title":"Multicolour\nChemical Imaging of Plant Tissues with Hyperspectral Stimulated Raman\nScattering Microscopy","volume":"146","author":"Iino","year":"2021","journal-title":"Analyst"},{"key":"2026083104104745000_cit24","doi-asserted-by":"publisher","first-page":"507","DOI":"10.1146\/annurev.physchem.012809.103512","article-title":"Coherent Nonlinear\nOptical Imaging: Beyond Fluorescence Microscopy","volume":"62","author":"Min","year":"2011","journal-title":"Annu. Rev. Phys. Chem."},{"key":"2026083104104745000_cit25","doi-asserted-by":"publisher","first-page":"1994","DOI":"10.1021\/acs.est.1c05818","article-title":"Label-Free Multiphoton\nImaging of Microbes in Root,\nMineral, and Soil Matrices with Time-Gated Coherent Raman and Fluorescence\nLifetime Imaging","volume":"56","author":"Lee","year":"2022","journal-title":"Environ. Sci. Technol."},{"key":"2026083104104745000_cit26","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1364\/OL.33.000923","article-title":"Ultrabroadband\nMultiplex CARS Microspectroscopy and Imaging Using a Subnanosecond\nSupercontinuum Light Source in the Deep near Infrared","volume":"33","author":"Okuno","year":"2008","journal-title":"Opt. Lett."},{"key":"2026083104104745000_cit27","doi-asserted-by":"publisher","first-page":"2708","DOI":"10.1021\/acs.jpcb.3c00302","article-title":"Simultaneous Imaging and Characterization\nof Polyunsaturated\nFatty Acids, Carotenoids, and Microcrystalline Guanine in Single Aurantiochytrium limacinum Cells with Linear and Nonlinear\nRaman Microspectroscopy","volume":"127","author":"Sasaki","year":"2023","journal-title":"J. Phys. Chem. B"},{"key":"2026083104104745000_cit28","doi-asserted-by":"publisher","first-page":"39956","DOI":"10.1021\/acsomega.4c05706","article-title":"Multimodal\nMolecular\nImaging Reveals a Novel Membrane Component in Sporangia of the Rare\nActinomycete Actinoplanes missouriensis","volume":"9","author":"Usami","year":"2024","journal-title":"ACS Omega"},{"key":"2026083104104745000_cit29","doi-asserted-by":"publisher","first-page":"3622","DOI":"10.1364\/OE.14.003622","article-title":"Direct Extraction of\nRaman Line-Shapes from Congested CARS Spectra","volume":"14","author":"Vartiainen","year":"2006","journal-title":"Opt. Express"},{"key":"2026083104104745000_cit30","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1002\/cphc.200600481","article-title":"Quantitative\nCARS Spectroscopy Using the Maximum Entropy Method: The Main Lipid\nPhase Transition","volume":"8","author":"Rinia","year":"2007","journal-title":"ChemPhysChem"},{"key":"2026083104104745000_cit31","doi-asserted-by":"publisher","first-page":"6773","DOI":"10.1002\/anie.201001560","article-title":"Quantitative CARS Molecular Fingerprinting\nof Single Living Cells with the Use of the Maximum Entropy Method","volume":"49","author":"Okuno","year":"2010","journal-title":"Angew. Chem., Int. Ed."},{"key":"2026083104104745000_cit32","doi-asserted-by":"publisher","first-page":"1174","DOI":"10.1002\/jrs.1747","article-title":"Noise Reduction in Raman Spectra:\nFinite Impulse Response Filtration Versus Savitzky\u2013Golay Smoothing","volume":"38","author":"C\u030clupek","year":"2007","journal-title":"J. Raman Spectrosc."},{"key":"2026083104104745000_cit33","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1081\/ASR-100106157","article-title":"FT-Infrared\nand FT-Raman Spectroscopy in Biomedical Research","volume":"36","author":"Naumann","year":"2001","journal-title":"Appl. Spectrosc. Rev."},{"key":"2026083104104745000_cit34","doi-asserted-by":"publisher","first-page":"10009","DOI":"10.1021\/bi050179w","article-title":"Molecular-Level\nInvestigation of the Structure, Transformation, and Bioactivity of\nSingle Living Fission Yeast Cells by Time- and Space-Resolved Raman\nSpectroscopy","volume":"44","author":"Huang","year":"2005","journal-title":"Biochemistry"},{"key":"2026083104104745000_cit35","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1002\/jrs.4607","article-title":"Raman Spectroscopy of Lipids: A Review","volume":"46","author":"Czamara","year":"2015","journal-title":"J. Raman Spectrosc."},{"key":"2026083104104745000_cit36","doi-asserted-by":"publisher","first-page":"1061","DOI":"10.1002\/jrs.4335","article-title":"Raman Spectroscopy\nof Proteins: A Review","volume":"44","author":"Rygula","year":"2013","journal-title":"J. Raman Spectrosc."},{"key":"2026083104104745000_cit37","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1104\/pp.010466","article-title":"Translocation\nand Utilization of Fungal Storage Lipid in the Arbuscular Mycorrhizal\nSymbiosis","volume":"128","author":"Bago","year":"2002","journal-title":"Plant Physiol."},{"key":"2026083104104745000_cit38","doi-asserted-by":"publisher","first-page":"578919","DOI":"10.3389\/fpls.2020.578919","article-title":"Responses of Arbuscular Mycorrhizal\nSymbiosis to Abiotic Stress: A Lipid-Centric Perspective","volume":"11","author":"Feng","year":"2020","journal-title":"Front. Plant Sci."},{"key":"2026083104104745000_cit39","doi-asserted-by":"publisher","DOI":"10.7554\/elife.29107","article-title":"Lipid Transfer from Plants to Arbuscular Mycorrhiza\nFungi","volume":"6","author":"Keymer","year":"2017","journal-title":"eLife"},{"key":"2026083104104745000_cit40","doi-asserted-by":"publisher","first-page":"1172","DOI":"10.1126\/science.aam9970","article-title":"Plants Transfer Lipids to Sustain Colonization by Mutualistic\nMycorrhizal and Parasitic Fungi","volume":"356","author":"Jiang","year":"2017","journal-title":"Science"},{"key":"2026083104104745000_cit41","doi-asserted-by":"publisher","first-page":"1175","DOI":"10.1126\/science.aan0081","article-title":"Fatty Acids\nin Arbuscular\nMycorrhizal Fungi Are Synthesized by the Host Plant","volume":"356","author":"Luginbuehl","year":"2017","journal-title":"Science"},{"key":"2026083104104745000_cit42","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1111\/j.1469-8137.1978.tb02274.x","article-title":"Lipid Physiology\nof Vesicular-Arbuscular Mycorrhiza","volume":"80","author":"Cooper","year":"1978","journal-title":"New Phytol."},{"key":"2026083104104745000_cit43","doi-asserted-by":"publisher","first-page":"6762","DOI":"10.1128\/aem.69.11.6762-6767.2003","article-title":"Fungal Lipid Accumulation and Development\nof Mycelial Structures by Two Arbuscular Mycorrhizal Fungi","volume":"69","author":"van Aarle","year":"2003","journal-title":"Appl. Environ. Microbiol."},{"key":"2026083104104745000_cit44","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/0008-6215(87)80306-3","article-title":"Band Assignments\nin the Raman Spectra of Celluloses","volume":"160","author":"Wiley","year":"1987","journal-title":"Carbohydr.\nRes."},{"key":"2026083104104745000_cit45","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1016\/j.aca.2011.08.031","article-title":"1064 nm\nDispersive Multichannel Raman\nSpectroscopy for the Analysis of Plant Lignin","volume":"706","author":"Meyer","year":"2011","journal-title":"Anal. Chim. Acta"},{"key":"2026083104104745000_cit46","doi-asserted-by":"publisher","first-page":"561","DOI":"10.1002\/jrs.1185","article-title":"Raman Spectroscopic Study of the\nEpicuticular Wax Layer of Mature Mango (Mangifera indica) Fruit","volume":"35","author":"Prinsloo","year":"2004","journal-title":"J. Raman Spectrosc."},{"key":"2026083104104745000_cit47","doi-asserted-by":"publisher","first-page":"2383","DOI":"10.1016\/S1386-1425(97)00178-9","article-title":"FT Raman Microscopy of Untreated\nNatural Plant Fibres","volume":"53","author":"Edwards","year":"1997","journal-title":"Spectrochim. Acta, Part\nA"},{"key":"2026083104104745000_cit48","doi-asserted-by":"publisher","first-page":"10101","DOI":"10.1073\/pnas.1205726109","article-title":"Casparian\nStrip Diffusion Barrier in Arabidopsis Is Made of\na Lignin Polymer without Suberin","volume":"109","author":"Naseer","year":"2012","journal-title":"Proc. Natl.\nAcad. Sci. U.S.A."},{"key":"2026083104104745000_cit49","doi-asserted-by":"publisher","first-page":"4142","DOI":"10.1103\/PhysRevLett.82.4142","article-title":"Three-Dimensional\nVibrational Imaging\nby Coherent Anti-Stokes Raman Scattering","volume":"82","author":"Zumbusch","year":"1999","journal-title":"Phys.\nRev. Lett."},{"key":"2026083104104745000_cit50","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1007\/s00572-022-01077-2","article-title":"Physiological\nand Transcriptomic Response of Medicago truncatula to Colonization by High- or Low-Benefit Arbuscular Mycorrhizal Fungi","volume":"32","author":"Cope","year":"2022","journal-title":"Mycorrhiza"},{"key":"2026083104104745000_cit51","doi-asserted-by":"publisher","first-page":"1206","DOI":"10.1016\/j.cub.2017.03.003","article-title":"A Transcriptional Program for Arbuscule Degeneration\nDuring AM Symbiosis Is Regulated by MYB1","volume":"27","author":"Floss","year":"2017","journal-title":"Curr.\nBiol."},{"key":"2026083104104745000_cit52","doi-asserted-by":"publisher","first-page":"1720","DOI":"10.1073\/pnas.0608136104","article-title":"A Medicago truncatula Phosphate Transporter Indispensable\nfor the Arbuscular Mycorrhizal Symbiosis","volume":"104","author":"Javot","year":"2007","journal-title":"Proc.\nNatl. Acad. Sci. U.S.A."},{"key":"2026083104104745000_cit53","doi-asserted-by":"publisher","DOI":"10.1002\/jbio.201960163","article-title":"Organelle Specific Simultaneous Raman\/Green\nFluorescence Protein Microspectroscopy for Living Cell Physicochemical\nStudies","volume":"13","author":"Wattanavichean","year":"2020","journal-title":"J. Biophotonics"},{"key":"2026083104104745000_cit54","doi-asserted-by":"publisher","first-page":"43569","DOI":"10.1038\/srep43569","article-title":"Protein Expression\nGuided Chemical Profiling of Living Cells by the Simultaneous Observation\nof Raman Scattering and Anti-Stokes Fluorescence Emission","volume":"7","author":"Chiu","year":"2017","journal-title":"Sci. Rep."},{"key":"2026083104104745000_cit55","doi-asserted-by":"publisher","first-page":"13324","DOI":"10.1073\/pnas.202474599","article-title":"Rice Phosphate Transporters\nInclude an Evolutionarily Divergent Gene Specifically Activated in\nArbuscular Mycorrhizal Symbiosis","volume":"99","author":"Paszkowski","year":"2002","journal-title":"Proc. Natl.\nAca. Sci. 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