{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T13:06:34Z","timestamp":1789477594041,"version":"build-2803163510"},"reference-count":47,"publisher":"American Chemical Society (ACS)","issue":"18","license":[{"start":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T00:00:00Z","timestamp":1787529600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T00:00:00Z","timestamp":1787529600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T00:00:00Z","timestamp":1787529600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["GM159321"],"award-info":[{"award-number":["GM159321"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,15]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>A small fraction of vascular plants has evolved the remarkable ability for their vegetative tissues to survive the loss of the vast majority of water and revive upon rehydration. Understanding the chemical and structural mechanisms for this desiccation tolerance is important for improving agricultural crop productivity in our changing climate. Although gas chromatography\u2013mass spectrometry has been used to profile metabolites in desiccation-tolerant plants, these studies involve chemical extraction and hence do not provide quantitative information about insoluble species in whole plants. Here, we employ solid-state NMR spectroscopy to examine the intact leaves and stems of Selaginella lepidophylla, a well-known resurrection plant, in the hydrated resurrected state and the dry dormant state. Using rigid-selective 13C cross-polarization (CP), dynamic-selective 13C insensitive-nuclei-enhancement-by-polarization transfer (INEPT), and direct polarization 13C NMR experiments, we show that the dormant state of S. lepidophylla contains higher concentrations of dynamic lipids, most likely triglycerides, as well as higher levels of immobilized trehalose and other carbohydrates compared to the hydrated state. Two control plants that are desiccation-sensitive lack the high level of dynamic lipids, and the lipids that are present have distinct chemical shifts from those of S. lepidophylla. These results indicate that dynamic lipids play a significant role in the adaptive response of vascular plants to severe water loss.<\/jats:p>","DOI":"10.1021\/acs.biochem.6c00400","type":"journal-article","created":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T12:07:08Z","timestamp":1789474028000},"page":"2875-2884","source":"Crossref","is-referenced-by-count":0,"title":["Structures and\nDynamics of Metabolites in Desiccation-Tolerant\n                    <i>S. lepidophylla<\/i>\n                    Plant Whole Cells Investigated by 13C Solid-State NMR"],"prefix":"10.1021","volume":"65","author":[{"ORCID":"https:\/\/orcid.org\/0009-0003-8848-4213","authenticated-orcid":true,"given":"Yanina","family":"Pankratova","sequence":"first","affiliation":[{"name":"Massachusetts Institute of Technology , , 170 Albany Street , , ,","place":["Cambridge, Massachusetts, United States, 02139"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5255-5858","authenticated-orcid":true,"given":"Mei","family":"Hong","sequence":"additional","affiliation":[{"name":"Massachusetts Institute of Technology , , 170 Albany Street , , ,","place":["Cambridge, Massachusetts, United States, 02139"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,24]]},"reference":[{"key":"2026091508070273800_cit1","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1038\/s41477-018-0330-7","article-title":"The molecular\nstructure of plant sporopollenin","volume":"5","author":"Li","year":"2019","journal-title":"Nat. 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Bot."},{"key":"2026091508070273800_cit5","doi-asserted-by":"publisher","first-page":"983","DOI":"10.1111\/tpj.12008","article-title":"Comparative metabolic\nprofiling between desiccation-sensitive and desiccation-tolerant species\nof Selaginella reveals insights into the resurrection trait","volume":"72","author":"Yobi","year":"2012","journal-title":"Plant J."},{"key":"2026091508070273800_cit6","doi-asserted-by":"publisher","first-page":"369","DOI":"10.1093\/mp\/sss155","article-title":"Metabolomic profiling\nin Selaginella lepidophylla at various hydration states provides new\ninsights into the mechanistic basis of desiccation tolerance","volume":"6","author":"Yobi","year":"2013","journal-title":"Mol. Plant"},{"key":"2026091508070273800_cit7","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/0305-1978(90)90044-G","article-title":"Comparison of free sugars in growing\nand desiccated plants of Selaginella lepidophylla","volume":"18","author":"Adams","year":"1990","journal-title":"Biochem. Syst. Ecol."},{"key":"2026091508070273800_cit8","doi-asserted-by":"publisher","DOI":"10.1016\/j.plantsci.2021.111129","article-title":"Overexpression of the Selaginella\nlepidophylla bHLH\ntranscription factor enhances water-use efficiency, growth, and development\nin Arabidopsis","volume":"315","author":"Ariyarathne","year":"2022","journal-title":"Plant Sci."},{"key":"2026091508070273800_cit9","doi-asserted-by":"publisher","first-page":"1173","DOI":"10.1016\/j.plantsci.2006.02.004","article-title":"An EST\ncatalogue from the resurrection\nplant Selaginella lepidophylla reveals abiotic stress-adaptive genes","volume":"170","author":"Iturriaga","year":"2006","journal-title":"Plant Sci."},{"key":"2026091508070273800_cit10","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-017-02546-5","article-title":"Extreme\nhaplotype variation in the desiccation-tolerant clubmoss Selaginella\nlepidophylla","volume":"9","author":"VanBuren","year":"2018","journal-title":"Nat. Commun."},{"key":"2026091508070273800_cit11","doi-asserted-by":"publisher","DOI":"10.1038\/srep08064","article-title":"Hydro-Responsive\nCurling\nof the Resurrection Plant Selaginella lepidophylla","volume":"5","author":"Rafsanjani","year":"2015","journal-title":"Sci. Rep."},{"key":"2026091508070273800_cit12","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1078\/0367-2530-00018","article-title":"Structural and ultrastructural\ncharacterization of Selaginella lepidophylla, a desiccation-tolerant\nplant, during the rehydration process","volume":"197","author":"Brighigna","year":"2002","journal-title":"Flora"},{"key":"2026091508070273800_cit13","doi-asserted-by":"publisher","DOI":"10.1038\/s43586-020-00002-1","article-title":"Solid-state NMR spectroscopy","volume":"1","author":"Reif","year":"2021","journal-title":"Nat. Rev. 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Phys."},{"key":"2026091508070273800_cit20","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1007\/s12013-020-00947-w","article-title":"Lipids Composition\nin Plant Membranes","volume":"78","author":"Reszczyn\u0301ska","year":"2020","journal-title":"Cell Biochem. Biophys."},{"key":"2026091508070273800_cit21","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1104\/pp.113.222737","article-title":"The Formation\nand Function of Plant Cuticles","volume":"163","author":"Yeats","year":"2013","journal-title":"Plant Physiol."},{"key":"2026091508070273800_cit22","doi-asserted-by":"publisher","first-page":"316","DOI":"10.1016\/j.jmr.2006.05.017","article-title":"Multi-dimensional\n1H\u201313C HETCOR and FSLG-HETCOR\nNMR study of sphingomyelin bilayers containing cholesterol in the\ngel and liquid crystalline states","volume":"181","author":"Holland","year":"2006","journal-title":"J. Magn.\nReson."},{"key":"2026091508070273800_cit23","doi-asserted-by":"publisher","first-page":"1939","DOI":"10.1016\/S0006-3495(95)80064-0","article-title":"Study of phospholipid structure by\n1H, 13C, and 31P dipolar couplings from 2D NMR","volume":"69","author":"Hong","year":"1995","journal-title":"Biophys. 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Sci."},{"key":"2026091508070273800_cit33","doi-asserted-by":"publisher","first-page":"19374","DOI":"10.1021\/jacs.1c07429","article-title":"Identification and\nQuantification of Glycans in Whole Cells: Architecture\nof Microalgal Polysaccharides Described by Solid-State Nuclear Magnetic\nResonance","volume":"143","author":"Poulhazan","year":"2021","journal-title":"J. Am. Chem. Soc."},{"key":"2026091508070273800_cit34","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1016\/S0163-7827(01)00013-3","article-title":"The biogenesis\nand functions of lipid bodies in animals, plants and microorganisms","volume":"40","author":"Murphy","year":"2001","journal-title":"Prog. Lipid Res."},{"key":"2026091508070273800_cit35","doi-asserted-by":"publisher","first-page":"983","DOI":"10.1016\/j.molp.2018.05.003","article-title":"Genome\nanalysis of the ancient tracheophyte Selaginella tamariscina reveals\nevolutionary features relevant to the acquisition of desiccation tolerance","volume":"11","author":"Xu","year":"2018","journal-title":"Mol. Plant"},{"key":"2026091508070273800_cit36","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1104\/pp.118.3.975","article-title":"Drying\nincreases intracellular partitioning\nof amphiphilic substances into the lipid phase. Impact On membrane\npermeability and significance for desiccation tolerance","volume":"118","author":"Golovina","year":"1998","journal-title":"Plant Physiol"},{"key":"2026091508070273800_cit37","doi-asserted-by":"publisher","first-page":"579","DOI":"10.1146\/annurev.ph.54.030192.003051","article-title":"Anhydrobiosis","volume":"54","author":"Crowe","year":"1992","journal-title":"Annu. Rev. Physiol."},{"key":"2026091508070273800_cit38","doi-asserted-by":"publisher","first-page":"431","DOI":"10.1016\/S1360-1385(01)02052-0","article-title":"Mechanisms\nof plant desiccation tolerance","volume":"6","author":"Hoekstra","year":"2001","journal-title":"Trends Plant\nSci."},{"key":"2026091508070273800_cit39","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.carres.2015.04.006","article-title":"Structural analysis of novel trehalose-based oligosaccharides\nfrom extremely stress-tolerant ascospores of Neosartorya fischeri\n(Aspergillus fischeri)","volume":"411","author":"Wyatt","year":"2015","journal-title":"Carbohydr. Res."},{"key":"2026091508070273800_cit40","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1016\/S0926-2040(98)00069-1","article-title":"Solid-state\n13C NMR investigations of the glycosidic\nlinkage in alpha-alpha\u2019 trehalose","volume":"12","author":"Zhang","year":"1998","journal-title":"Solid\nState Nucl. Magn. Reson."},{"key":"2026091508070273800_cit41","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1016\/0008-6215(93)84096-O","article-title":"(13C)-substituted sucrose: 13C-1H\nand 13C-13C spin\ncoupling constants to assess furanose ring and glycosidic bond conformations\nin aqueous solution","volume":"249","author":"Duker","year":"1993","journal-title":"Carbohydr. Res."},{"key":"2026091508070273800_cit42","doi-asserted-by":"publisher","DOI":"10.1016\/j.foodchem.2022.133125","article-title":"Quantification\nof polyols in sugar-free foodstuffs\nby qNMR","volume":"390","author":"Scettri","year":"2022","journal-title":"Food Chem."},{"key":"2026091508070273800_cit43","doi-asserted-by":"publisher","DOI":"10.3390\/molecules28031363","article-title":"Byproducts of Globe Artichoke and\nCauliflower Production as a New Source of Bioactive Compounds in the\nGreen Economy Perspective: An NMR Study","volume":"28","author":"Ingallina","year":"2023","journal-title":"Molecules"},{"key":"2026091508070273800_cit44","doi-asserted-by":"publisher","DOI":"10.3390\/molecules31040662","article-title":"Phytohemical Profiling, Bioactivity and Toxicity Evaluation\nof Elsholtzia cypriani, a Potential Multifunctional Natural Feed Additive","volume":"31","author":"Zhang","year":"2026","journal-title":"Molecules"},{"key":"2026091508070273800_cit45","doi-asserted-by":"publisher","first-page":"763","DOI":"10.1023\/A:1015810303089","article-title":"One-dimensional\nand two-dimensional 1H- and 13C-nuclear\nmagnetic resonance (NMR) analysis of vitamin E raw materials or analytical\nreference standards","volume":"8","author":"Baker","year":"1991","journal-title":"Pharm. Res."},{"key":"2026091508070273800_cit46","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1351\/pac197647020097","article-title":"Carbon-13\nNMR Spectra of Carotenoids","volume":"47","author":"Moss","year":"1976","journal-title":"Pure Appl. Chem."},{"key":"2026091508070273800_cit47","doi-asserted-by":"publisher","first-page":"801","DOI":"10.1351\/pac198557050801","article-title":"NMR OF carotenoids\n- new experimental techniques","volume":"57","author":"Englert","year":"1985","journal-title":"Pure Appl. 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