{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T04:49:41Z","timestamp":1787028581620,"version":"build-2736575974"},"reference-count":24,"publisher":"MDPI AG","issue":"10","license":[{"start":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T00:00:00Z","timestamp":1779148800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82274055"],"award-info":[{"award-number":["82274055"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["82274055"],"award-info":[{"award-number":["82274055"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004608","name":"Natural Science Foundation of Jiangsu Province","doi-asserted-by":"publisher","award":["BK20250766"],"award-info":[{"award-number":["BK20250766"]}],"id":[{"id":"10.13039\/501100004608","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Open Project of Chinese Materia Medica First-Class Discipline of Nanjing University of Chinese Medicine","award":["ZYXYL2024-002"],"award-info":[{"award-number":["ZYXYL2024-002"]}]},{"name":"Wu Qinan National TCM Herbalist Heritage Workshop Research Grant","award":["2025"],"award-info":[{"award-number":["2025"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Sparganium stoloniferum tubers (SL), known medicinally as Sparganii Rhizoma, are commonly considered superior at the winter-harvest stage, when they show the traditional quality traits of heavy weight and firm texture. However, the developmental basis of this quality phenotype remains insufficiently understood. This study aimed to determine how tissue organization, cell-wall architecture, starch deposition, and related transcriptional patterns are associated with winter-harvest quality in SL. By comparing SL at different developmental stages, we found that maturation was accompanied by reduced moisture content, increased tuber density, higher parenchyma cell density, progressive cell-wall thickening, and marked starch accumulation. Laser scanning confocal microscopy (LSCM), scanning electron microscopy (SEM), and transmission electron microscopy (TEM) observations further revealed thickened multilamellar cell walls and abundant clustered or compound-like starch bodies in mature SL. Starch isolated from mature SL displayed an A-type crystalline pattern, short-range order, and high gelatinization and pasting temperatures, indicating an ordered and thermally stable starch matrix. Cell-wall Fourier-transform infrared spectroscopy (FTIR) and solid-state nuclear magnetic resonance (NMR) analyses showed a predominantly polysaccharide-rich framework with subtle maturation-associated changes in aromatic- and methoxy-associated wall signals. Transcript-guided pathway analysis, supported by reverse transcription quantitative polymerase chain reaction (RT\u2013qPCR)validation, suggested developmental shifts in carbohydrate metabolism, lipid-related metabolism, and gibberellin-associated transcriptional patterns. Together, these findings indicate that winter-harvest quality in SL is associated with coordinated tissue consolidation, cell-wall maturation, starch deposition, and transcriptional reprogramming, providing a structural and molecular framework for understanding the traditional firm-texture trait of S. stoloniferum.<\/jats:p>","DOI":"10.3390\/ijms27104566","type":"journal-article","created":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T15:17:23Z","timestamp":1779203843000},"page":"4566","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Coordinated Cell-Wall and Starch Maturation Is Associated with Winter-Harvest Quality in Sparganium stoloniferum Tubers"],"prefix":"10.3390","volume":"27","author":[{"given":"Xilong","family":"Qian","sequence":"first","affiliation":[{"name":"National Key Laboratory on Technologies for Chinese Medicine Pharmaceutical Process Control and Intelligent Manufacture, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing 210023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maoqi","family":"Pan","sequence":"additional","affiliation":[{"name":"School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing 210023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingying","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing 210023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinan","family":"Liu","sequence":"additional","affiliation":[{"name":"Nanjing Institute for Food and Drug Control, Nanjing 211198, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fan","family":"Yang","sequence":"additional","affiliation":[{"name":"National Key Laboratory on Technologies for Chinese Medicine Pharmaceutical Process Control and Intelligent Manufacture, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing 210023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chanchan","family":"Liu","sequence":"additional","affiliation":[{"name":"National Key Laboratory on Technologies for Chinese Medicine Pharmaceutical Process Control and Intelligent Manufacture, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing 210023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0543-1091","authenticated-orcid":false,"given":"Mengru","family":"Sang","sequence":"additional","affiliation":[{"name":"National Key Laboratory on Technologies for Chinese Medicine Pharmaceutical Process Control and Intelligent Manufacture, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing 210023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinan","family":"Wu","sequence":"additional","affiliation":[{"name":"National Key Laboratory on Technologies for Chinese Medicine Pharmaceutical Process Control and Intelligent Manufacture, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"Jiangsu Collaborative Innovation Center of Chinese Medicinal Resources Industrialization, Nanjing University of Chinese Medicine, Nanjing 210023, China"},{"name":"School of Pharmacy, Nanjing University of Chinese Medicine, Nanjing 210023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,5,19]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Pan, K., Lu, C., Nie, P., Hu, M., Zhou, X., Chen, X., and Wang, W. (2021). Predominantly Symplastic Phloem Unloading of Photosynthates Maintains Efficient Starch Accumulation in the Cassava Storage Roots (Manihot esculenta Crantz). BMC Plant Biol., 21.","DOI":"10.1186\/s12870-021-03088-1"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"7260","DOI":"10.1038\/s41467-024-51727-6","article-title":"Source-Sink Synergy Is the Key Unlocking Sweet Potato Starch Yield Potential","volume":"15","author":"Jiang","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Xu, L., Wang, J., Lei, M., Li, L., Fu, Y., Wang, Z., Ao, M., and Li, Z. (2016). Transcriptome Analysis of Storage Roots and Fibrous Roots of the Traditional Medicinal Herb Callerya speciosa (Champ.) Schot. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0160338"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1868","DOI":"10.1111\/1755-0998.13850","article-title":"Genomic Analysis of the Emergent Aquatic Plant Sparganium stoloniferum Provides Insights into Its Clonality, Local Adaptation and Demographic History","volume":"23","author":"Zou","year":"2023","journal-title":"Mol. Ecol. Resour."},{"key":"ref_5","unstructured":"Royal Botanic Gardens. K (2024, March 20). Plants of the World Online: Sparganium stoloniferum, Available online: http:\/\/www.plantsoftheworldonline.org."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"113571","DOI":"10.1016\/j.jep.2020.113571","article-title":"Sparganii Rhizoma: A Review of Traditional Clinical Application, Processing, Phytochemistry, Pharmacology, and Toxicity","volume":"268","author":"Jia","year":"2021","journal-title":"J. Ethnopharmacol."},{"key":"ref_7","unstructured":"Commission, C.P. (2025). Pharmacopoeia of the People\u2019s Republic of China: Volume I, China Medical Science and Technology Press."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"100400","DOI":"10.1016\/j.stress.2024.100400","article-title":"Multi-Omics Reveals Phenol-Based Coordinated Defense of Sparganium stoloniferum rhizoma","volume":"11","author":"Sang","year":"2024","journal-title":"Plant Stress"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"2769","DOI":"10.1093\/jxb\/ern142","article-title":"Antisense Inhibition of a Pectate Lyase Gene Supports a Role for Pectin Depolymerization in Strawberry Fruit Softening","volume":"59","author":"Matas","year":"2008","journal-title":"J. Exp. Bot."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Atkinson, R.G., Sutherland, P.W., Johnston, S.L., Gunaseelan, K., Hallett, I.C., Mitra, D., Brummell, D.A., Schr\u00f6der, R., Johnston, J.W., and Schaffer, R.J. (2012). Down-Regulation of POLYGALACTURONASE1 Alters Firmness, Tensile Strength and Water Loss in Apple (Malus \u00d7 domestica) Fruit. BMC Plant Biol., 12.","DOI":"10.1186\/1471-2229-12-129"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1038\/s41538-023-00230-1","article-title":"Lipid Complexation Reduces Rice Starch Digestibility and Boosts Short-Chain Fatty Acid Production Via Gut Microbiota","volume":"7","author":"Shen","year":"2023","journal-title":"NPJ Sci. Food"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"123530","DOI":"10.1016\/j.carbpol.2025.123530","article-title":"Amylose Content Controls the Structures and Digestibility of Starch-Lipid Complexes During Microwave Postprocessing","volume":"358","author":"Tan","year":"2025","journal-title":"Carbohydr. Polym."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"103050","DOI":"10.1016\/j.fochx.2025.103050","article-title":"Effects of Oat Lipid-Starch Complexation on Starch Structure, Freeze-Thaw Stability and Pasting Characteristics Under Ultra-High Pressure Treatment","volume":"31","author":"Xie","year":"2025","journal-title":"Food Chem. X"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1038\/s41438-021-00589-w","article-title":"Antagonism Between Abscisic Acid and Gibberellin Regulates Starch Synthesis and Corm Development in Gladiolus hybridus","volume":"8","author":"Li","year":"2021","journal-title":"Hortic. Res."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Meng, G., Yong, M., Zhang, Z., Zhang, Y., Wang, Y., Xiong, A., and Su, X. (2024). Exogenous Gibberellin Suppressed Taproot Secondary Thickening by Inhibiting the Formation and Maintenance of Vascular Cambium in Radish (Raphanus sativus L.). Front. Plant Sci., 15.","DOI":"10.3389\/fpls.2024.1395999"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1365-313X.1993.tb00007.x","article-title":"Structural Models of Primary Cell Walls in Flowering Plants: Consistency of Molecular Structure with the Physical Properties of the Walls During Growth","volume":"3","author":"Carpita","year":"1993","journal-title":"Plant J."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1104\/pp.110.161240","article-title":"Plant Cell Walls","volume":"154","author":"Keegstra","year":"2010","journal-title":"Plant Physiol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1257","DOI":"10.1093\/plcell\/koad325","article-title":"The Plant Cell Wall\u2014Dynamic, Strong, and Adaptable\u2014Is a Natural Shapeshifter","volume":"36","author":"Delmer","year":"2024","journal-title":"Plant Cell"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1046\/j.1365-3040.2003.01034.x","article-title":"Intercellular Adhesion and Cell Separation in Plants","volume":"26","author":"Jarvis","year":"2003","journal-title":"Plant Cell Environ."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1002\/1521-379X(200104)53:3\/4<121::AID-STAR121>3.0.CO;2-Q","article-title":"Relevance of Amylose-Lipid Complexes to the Behaviour of Thermally Processed Starches","volume":"53","author":"Becker","year":"2001","journal-title":"Starch St\u00e4rke"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"143450","DOI":"10.1016\/j.foodchem.2025.143450","article-title":"Preparation and Functional Characteristics of Starch-Lipid Complexes with Different Oleic Acid-Rich Glycerolipids","volume":"476","author":"Chen","year":"2025","journal-title":"Food Chem."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Hong, C.P., Kim, J., Lee, J., Yoo, S.I., Bae, W., Geem, K.R., Yu, J., Jang, I., Jo, I.H., and Cho, H. (2021). Gibberellin Signaling Promotes the Secondary Growth of Storage Roots in Panax ginseng. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22168694"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Singh, V., Sergeeva, L., Ligterink, W., Aloni, R., Zemach, H., Doron-Faigenboim, A., Yang, J., Zhang, P., Shabtai, S., and Firon, N. (2019). Gibberellin Promotes Sweetpotato Root Vascular Lignification and Reduces Storage-Root Formation. Front Plant Sci., 10.","DOI":"10.3389\/fpls.2019.01320"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of Relative Gene Expression Data Using Real-Time Quantitative PCR and the 2\u2212\u0394\u0394Ct Method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/10\/4566\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,25]],"date-time":"2026-05-25T04:18:35Z","timestamp":1779682715000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/10\/4566"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,19]]},"references-count":24,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2026,5]]}},"alternative-id":["ijms27104566"],"URL":"https:\/\/doi.org\/10.3390\/ijms27104566","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,19]]}}}