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We examined the transition from a cutin-dominated cuticle in common cucumber to a ligno-suberized periderm in Sikkim cucumber. Multiscale microscopy and histochemistry resolved a developmental shift from a smooth epidermis to a multilayered phellem characterized by distinct suberin lamellae and lignin deposition. GC-MS-based profiling revealed that periderm formation involves a shift from C16\/C18 cutin monomers to an enrichment in &amp;gt;C20 very-long-chain aliphatics (fatty acids, alcohols, hydroxy acids) and phenolics like ferulic acid, hydroxybenzoic acids, and p-hydroxyphenyl, guaiacyl, and syringyl lignin moieties. ssNMR confirmed the increase in oxygenated and phenolic-rich polymer assemblies, consistent with higher cross-link density. Accordingly, peridermal surfaces exhibited significantly enhanced stiffness, adhesion, and elastic resistance. DSC revealed significant elevations in glass transition temperatures (Tg) from 11 \u00b0C to 60 \u00b0C. These findings establish the fruit periderm as a mechanically robust, thermally stable composite model for sustainable biopolymer design.<\/jats:p>","DOI":"10.1021\/acs.biomac.6c00879","type":"journal-article","created":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T15:35:56Z","timestamp":1783352156000},"page":"5570-5580","source":"Crossref","is-referenced-by-count":1,"title":["Ligno-Suberized\nFruit Periderm as a Mechanically Reinforced\nBiomacromolecular Composite"],"prefix":"10.1021","volume":"27","author":[{"given":"Yaarit","family":"Kutsher","sequence":"first","affiliation":[{"name":"Volcani Institute , , ,","place":["Rishon LeZion, Israel, 7505101"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ekram","family":"Wassel","sequence":"additional","affiliation":[{"name":"The City College of New York (CCNY), City University of New York (CUNY), and Institute for Macromolecular Assemblies , , , ,","place":["New York, New York, United States, 10031"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Austin","family":"Laviano","sequence":"additional","affiliation":[{"name":"The City College of New York (CCNY), City University of New York (CUNY), and Institute for Macromolecular Assemblies , , , ,","place":["New York, New York, United States, 10031"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jorge","family":"Carvajal","sequence":"additional","affiliation":[{"name":"University of Bonn , , Kirschallee 1 , ,","place":["Bonn, Germany, 53115"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ekaterina","family":"Manasherova","sequence":"additional","affiliation":[{"name":"Volcani Institute , , ,","place":["Rishon LeZion, Israel, 7505101"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Viktoria V.","family":"Zeisler-Diehl","sequence":"additional","affiliation":[{"name":"University of Bonn , , Kirschallee 1 , ,","place":["Bonn, Germany, 53115"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7003-9929","authenticated-orcid":true,"given":"Lukas","family":"Schreiber","sequence":"additional","affiliation":[{"name":"University of Bonn , , Kirschallee 1 , ,","place":["Bonn, Germany, 53115"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruth E.","family":"Stark","sequence":"additional","affiliation":[{"name":"The City College of New York (CCNY), City University of New York (CUNY), and Institute for Macromolecular Assemblies , , , ,","place":["New York, New York, United States, 10031"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4209-0257","authenticated-orcid":true,"given":"Hagai","family":"Cohen","sequence":"additional","affiliation":[{"name":"Volcani Institute , , ,","place":["Rishon LeZion, Israel, 7505101"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,7,6]]},"reference":[{"key":"2026081008082741500_cit1","doi-asserted-by":"publisher","first-page":"4639","DOI":"10.1093\/jxb\/eru301","article-title":"There\u2019s more than one way to skin a fruit: formation\nand functions of fruit cuticles","volume":"65","author":"Martin","year":"2013","journal-title":"J. Exp. Bot."},{"key":"2026081008082741500_cit2","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1016\/j.postharvbio.2013.08.012","article-title":"The fruit\ncuticle as a modulator\nof postharvest quality","volume":"87","author":"Lara","year":"2014","journal-title":"Postharvest Biol. Technol."},{"key":"2026081008082741500_cit3","doi-asserted-by":"publisher","first-page":"770","DOI":"10.3389\/fpls.2019.00770","article-title":"Shelf life\npotential and the fruit\ncuticle: the unexpected player","volume":"10","author":"Lara","year":"2019","journal-title":"Front. Plant\nSci."},{"key":"2026081008082741500_cit4","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2021.663165","article-title":"The plant\ncuticle: an ancient guardian barrier set against long-standing rivals","volume":"12","author":"Arya","year":"2021","journal-title":"Front. Plant Sci."},{"issue":"2","key":"2026081008082741500_cit5","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1111\/j.1365-313X.2009.03969.x","article-title":"Cutin deficiency in the tomato fruit\ncuticle consistently\naffects resistance to microbial infection and biomechanical properties,\nbut not transpirational water loss","volume":"60","author":"Isaacson","year":"2009","journal-title":"Plant J."},{"issue":"1","key":"2026081008082741500_cit6","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1104\/pp.113.222737","article-title":"The formation and function of plant cuticles","volume":"163","author":"Yeats","year":"2013","journal-title":"Plant Physiol."},{"key":"2026081008082741500_cit7","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1002\/9780470988718.ch4","article-title":"Composition of plant cuticular\nwaxes","volume-title":"Biology of the Plant Cuticle","author":"Jetter","year":"2006"},{"key":"2026081008082741500_cit8","doi-asserted-by":"publisher","DOI":"10.1093\/aobpla\/pls048","article-title":"Russeting\nin apple and pear: a plastic\nperiderm replaces a stiff cuticle","volume":"5","author":"Khanal","year":"2013","journal-title":"AoB Plants."},{"issue":"21","key":"2026081008082741500_cit9","doi-asserted-by":"publisher","first-page":"6579","DOI":"10.1093\/jxb\/erv366","article-title":"Genome investigation\nsuggests MdSHN3,\nan APETALA2-domain transcription factor gene, to be a positive regulator\nof apple fruit cuticle formation and an inhibitor of russet development","volume":"66","author":"Lashbrooke","year":"2015","journal-title":"J. Exp. Bot."},{"issue":"4","key":"2026081008082741500_cit10","doi-asserted-by":"publisher","first-page":"1486","DOI":"10.1104\/pp.18.01158","article-title":"A multilevel study\nof melon fruit\nreticulation provides insight into skin ligno-suberization hallmarks","volume":"179","author":"Cohen","year":"2019","journal-title":"Plant Physiol."},{"key":"2026081008082741500_cit11","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1146\/annurev-arplant-102720-031405","article-title":"The making of plant armor: the periderm","volume":"73","author":"Serra","year":"2022","journal-title":"Annu.\nRev. Plant Biol."},{"key":"2026081008082741500_cit12","doi-asserted-by":"publisher","first-page":"190","DOI":"10.1080\/07352689.2017.1369333","article-title":"Ongoing growth challenges\nfruit skin integrity","volume":"36","author":"Knoche","year":"2017","journal-title":"Crit. Rev. 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Plant\nBiol."},{"issue":"2","key":"2026081008082741500_cit15","doi-asserted-by":"publisher","first-page":"kiae653","DOI":"10.1093\/plphys\/kiae653","article-title":"Update on the structure\nand regulated\nbiosynthesis of the apoplastic polymers cutin and suberin","volume":"197","author":"Kosma","year":"2025","journal-title":"Plant Physiol."},{"key":"2026081008082741500_cit16","doi-asserted-by":"publisher","first-page":"519","DOI":"10.1146\/annurev.arplant.54.031902.134938","article-title":"Lignin biosynthesis","volume":"54","author":"Boerjan","year":"2003","journal-title":"Annu. Rev. Plant Biol."},{"key":"2026081008082741500_cit17","doi-asserted-by":"publisher","DOI":"10.1199\/tab.0152","article-title":"The phenylpropanoid\npathway in Arabidopsis","volume":"9","author":"Fraser","year":"2011","journal-title":"Arabidopsis Book"},{"issue":"12","key":"2026081008082741500_cit18","doi-asserted-by":"publisher","first-page":"2918","DOI":"10.1139\/b84-391","article-title":"Biochemistry\nand function of cutin and suberin","volume":"62","author":"Kolattukudy","year":"1984","journal-title":"Can. J. Bot."},{"issue":"8","key":"2026081008082741500_cit19","doi-asserted-by":"publisher","first-page":"2600","DOI":"10.1021\/ma9714880","article-title":"A WISE NMR approach\nto heterogeneous biopolymer mixtures:\ndynamics and domains in wounded potato tissues","volume":"31","author":"Yan","year":"1998","journal-title":"Macromolecules"},{"key":"2026081008082741500_cit20","doi-asserted-by":"publisher","DOI":"10.1093\/hr\/uhac092","article-title":"The metabolic and proteomic repertoires\nof periderm tissue in skin of the reticulated Sikkim cucumber fruit","volume":"9","author":"Arya","year":"2022","journal-title":"Hortic. Res."},{"key":"2026081008082741500_cit21","doi-asserted-by":"publisher","DOI":"10.1016\/j.postharvbio.2022.112071","article-title":"Cucumber fruit skin\nreticulation affects post-harvest traits","volume":"194","author":"Nomberg","year":"2022","journal-title":"Postharvest\nBiol. Techno."},{"issue":"4","key":"2026081008082741500_cit22","doi-asserted-by":"publisher","DOI":"10.1093\/hr\/uhad036","article-title":"Microscopic and metabolic\ninvestigations disclose the factors that lead to skin cracking in\nchili-type pepper fruit varieties","volume":"10","author":"Marinov","year":"2023","journal-title":"Hortic. Res."},{"key":"2026081008082741500_cit23","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodchem.2025.145837","article-title":"Postharvest\nstorage affects the chemical composition of the avocado fruit cuticle\nin a cultivar-specific manner","volume":"493","author":"Manasherova","year":"2025","journal-title":"Food Chem."},{"issue":"1","key":"2026081008082741500_cit24","doi-asserted-by":"publisher","DOI":"10.1093\/plphys\/kiaf376","article-title":"Exocarp-specific\nexpression of a fungal cutinase in tomato fruits alters cuticle ultrastructure,\nchemistry and nanomechanics","volume":"199","author":"Arya","year":"2025","journal-title":"Plant Physiol."},{"issue":"2","key":"2026081008082741500_cit25","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1080\/02773818508085193","article-title":"Thioacidolysis of lignin: comparison\nwith acidolysis","volume":"5","author":"Lapierre","year":"1985","journal-title":"J. Wood Chem. Technol."},{"issue":"2","key":"2026081008082741500_cit26","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1002\/mas.10072","article-title":"Mass spectrometry\nin the biosynthetic and structural investigation of lignins","volume":"23","author":"Reale","year":"2004","journal-title":"Mass Spectrom. Rev."},{"key":"2026081008082741500_cit27","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1007\/978-3-642-74065-7_23","article-title":"Thioacidolysis","volume-title":"Methods in\nLignin Chemistry","author":"Rolando","year":"1992"},{"issue":"1","key":"2026081008082741500_cit28","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1021\/acs.biomac.5b01321","article-title":"Solid-state 13C\nNMR delineates the architectural design\nof biopolymers in native and genetically altered tomato fruit cuticles","volume":"17","author":"Chatterjee","year":"2016","journal-title":"Biomacromolecules."},{"issue":"30","key":"2026081008082741500_cit29","doi-asserted-by":"publisher","first-page":"6810","DOI":"10.1021\/acs.jafc.5b03206","article-title":"Defensive\narmor of potato tubers:\nnonpolar metabolite profiling, antioxidant assessment, and solid-state\nNMR compositional analysis of suberin-enriched wound-healing tissues","volume":"63","author":"Dastmalchi","year":"2015","journal-title":"J. Agric. Food Chem."},{"key":"2026081008082741500_cit30","doi-asserted-by":"crossref","first-page":"580","DOI":"10.1533\/9781845690830.3.580","article-title":"Surface and\ninterfacial studies of plant biopolymers","volume-title":"Molecular Interfacial Phenomena of Polymers and Biopolymers","author":"Batteas","year":"2005"},{"issue":"9","key":"2026081008082741500_cit31","doi-asserted-by":"publisher","first-page":"12175","DOI":"10.3390\/s130912175","article-title":"Determination of the\nelastic properties of tomato fruit\ncells with an atomic force microscope","volume":"13","author":"Zdunek","year":"2013","journal-title":"Sensors."},{"issue":"5","key":"2026081008082741500_cit32","doi-asserted-by":"publisher","first-page":"2033","DOI":"10.1111\/nph.18862","article-title":"Cuticle architecture\nand mechanical properties: a functional relationship\ndelineated through correlated multimodal imaging","volume":"238","author":"Reynoud","year":"2023","journal-title":"New Phytol."},{"issue":"5","key":"2026081008082741500_cit33","doi-asserted-by":"publisher","first-page":"2761","DOI":"10.1016\/S0006-3495(00)76515-5","article-title":"The influence\nof water on the nanomechanical behavior of the plant biopolyester\ncutin as studied by AFM and solid-state NMR","volume":"79","author":"Round","year":"2000","journal-title":"Biophys. J."},{"key":"2026081008082741500_cit34","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2021.807723","article-title":"The response\nof tomato fruit cuticle\nmembranes against heat and light","volume":"12","author":"Beni\u0301tez","year":"2022","journal-title":"Front. Plant\nSci."},{"issue":"3","key":"2026081008082741500_cit35","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1016\/j.pbi.2007.04.004","article-title":"Suberin -\na biopolyester forming apoplastic plant interfaces","volume":"10","author":"Franke","year":"2007","journal-title":"Curr. Opin. 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Agric. Food Chem."},{"key":"2026081008082741500_cit40","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2025.145570","article-title":"Elucidating the modes\nof incorporation of the ferulic acid amides feruloyltyramine and feruloyloctopamine\ninto the lignin-suberin fraction of potato periderms","volume":"319","author":"Del Ri\u0301o","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"2026081008082741500_cit41","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1021\/acs.jafc.7b04638","article-title":"Variability\nin lignin composition and structure in cell walls of different parts\nof Macau\u0301ba (Acrocomia aculeata) palm fruit","volume":"66","author":"Rencoret","year":"2018","journal-title":"J. Agric. Food Chem."},{"key":"2026081008082741500_cit42","doi-asserted-by":"publisher","first-page":"2366","DOI":"10.1038\/s41598-025-86563-1","article-title":"Naturally russeted\nand wound russeted skins of mango\n(cv. \u2018Apple\u2019) show no differences in anatomy, chemical\ncomposition or gene expression","volume":"15","author":"Straube","year":"2025","journal-title":"Sci. Rep."},{"issue":"9","key":"2026081008082741500_cit43","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0274733","article-title":"Apple fruit periderms\n(russeting) induced by wounding or by moisture have the same histologies,\nchemistries and gene expressions","volume":"17","author":"Chen","year":"2022","journal-title":"PLoS One"},{"issue":"1","key":"2026081008082741500_cit44","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/j.postharvbio.2011.10.004","article-title":"Structural\nand physiological changes associated with the skin spot disorder in\napple","volume":"64","author":"Grimm","year":"2012","journal-title":"Postharvest Biol. Technol."},{"key":"2026081008082741500_cit45","doi-asserted-by":"publisher","DOI":"10.1016\/j.postharvbio.2023.112639","article-title":"The polyphenolic suberized\nperidermal skin of the Sikkim cucumber\nacts as an effective chemical barrier against Botrytis cinerea infection","volume":"208","author":"Nomberg","year":"2024","journal-title":"Postharvest Biol. Technol."}],"container-title":["Biomacromolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/acs.biomac.6c00879","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/bomaf6\/article-pdf\/27\/8\/5570\/65794168\/acs.biomac.6c00879.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/bomaf6\/article-pdf\/27\/8\/5570\/65794168\/acs.biomac.6c00879.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T12:24:27Z","timestamp":1786364667000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/bomaf6\/article\/27\/8\/5570\/5201444\/Ligno-Suberized-Fruit-Periderm-as-a-Mechanically"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,6]]},"references-count":45,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,7,6]]},"published-print":{"date-parts":[[2026,8,10]]}},"URL":"https:\/\/doi.org\/10.1021\/acs.biomac.6c00879","relation":{},"ISSN":["1525-7797","1526-4602"],"issn-type":[{"value":"1525-7797","type":"print"},{"value":"1526-4602","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,6]]}}}