{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,26]],"date-time":"2026-03-26T03:56:19Z","timestamp":1774497379946,"version":"3.50.1"},"reference-count":61,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2021,6,22]],"date-time":"2021-06-22T00:00:00Z","timestamp":1624320000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001319","name":"University of Westminster","doi-asserted-by":"publisher","award":["N\/A"],"award-info":[{"award-number":["N\/A"]}],"id":[{"id":"10.13039\/501100001319","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>The extensibility of synthetic polymers is routinely modulated by the addition of lower molecular weight spacing molecules known as plasticizers, and there is some evidence that water may have similar effects on plant cell walls. Furthermore, it appears that changes in wall hydration could affect wall behavior to a degree that seems likely to have physiological consequences at water potentials that many plants would experience under field conditions. Osmotica large enough to be excluded from plant cell walls and bacterial cellulose composites with other cell wall polysaccharides were used to alter their water content and to demonstrate that the relationship between water potential and degree of hydration of these materials is affected by their composition. Additionally, it was found that expansins facilitate rehydration of bacterial cellulose and cellulose composites and cause swelling of plant cell wall fragments in suspension and that these responses are also affected by polysaccharide composition. Given these observations, it seems probable that plant environmental responses include measures to regulate cell wall water content or mitigate the consequences of changes in wall hydration and that it may be possible to exploit such mechanisms to improve crop resilience.<\/jats:p>","DOI":"10.3390\/plants10071263","type":"journal-article","created":{"date-parts":[[2021,6,22]],"date-time":"2021-06-22T22:10:59Z","timestamp":1624399859000},"page":"1263","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":20,"title":["Plant Cell Wall Hydration and Plant Physiology: An Exploration of the Consequences of Direct Effects of Water Deficit on the Plant Cell Wall"],"prefix":"10.3390","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8251-6681","authenticated-orcid":false,"given":"David Stuart","family":"Thompson","sequence":"first","affiliation":[{"name":"School of Life Sciences, University of Westminster, 115 New Cavendish Street, London W1W 6UW, UK"}]},{"given":"Azharul","family":"Islam","sequence":"additional","affiliation":[{"name":"School of Life Sciences, University of Westminster, 115 New Cavendish Street, London W1W 6UW, UK"}]}],"member":"1968","published-online":{"date-parts":[[2021,6,22]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"388","DOI":"10.1111\/nph.12473","article-title":"My body is a cage: Mechanisms and modulation of plant cell growth","volume":"201","author":"Braidwood","year":"2014","journal-title":"New Phytol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"3223","DOI":"10.1093\/jxb\/ery181","article-title":"Turgor maintenance by osmotic adjustment: 40 years of progress","volume":"69","author":"Turner","year":"2018","journal-title":"J. Exp. Bot."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1111\/j.1399-3054.1997.tb04787.x","article-title":"Multiple signals and mechanisms that regulate leaf growth and stomatal behaviour during water deficit","volume":"100","author":"Thompson","year":"1997","journal-title":"Physiol. Plant."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1093\/jxb\/40.10.1099","article-title":"Derivation of plant cell wall water content by examination of the water-holding capacity of membrane-disrupted tissues","volume":"219","author":"Hardegree","year":"1989","journal-title":"J. Exp. Bot."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Fong, R.J., Robertson, A., Mallon, P.E., and Thompson, R.L. (2018). The impact of plasticizer and degree of hydrolysis on free volume of poly(vinyl alcohol) films. Polymers, 10.","DOI":"10.3390\/polym10091036"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1317","DOI":"10.1016\/j.carres.2004.02.027","article-title":"Material properties of concentrated pectin networks","volume":"339","author":"Zsivanovits","year":"2004","journal-title":"Carbohyd. Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1111\/j.1438-8677.1995.tb00508.x","article-title":"Water potential modulates extensibility of rye coleoptile cell walls","volume":"108","author":"Edelmann","year":"1995","journal-title":"Bot. Acta"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"3361","DOI":"10.1093\/jxb\/erm183","article-title":"Cell wall water content has a direct effect on extensibility in growing hypocotyls of sunflower (Helianthus annuus L.)","volume":"58","author":"Evered","year":"2007","journal-title":"J. Exp. Bot."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1021\/bm005632s","article-title":"Swelling behavior of the tomato cell wall network","volume":"2","author":"MacDougall","year":"2001","journal-title":"Biomacromolecules"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1675","DOI":"10.1093\/jxb\/45.Special_Issue.1675","article-title":"The physical chemistry of the primary cell wall: Implications for the control of expansion rate","volume":"45","author":"Passioura","year":"1994","journal-title":"J. Exp. Bot."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/j.plaphy.2017.11.005","article-title":"Pectin and cellulose cell wall composition enables different strategies to leaf water uptake in plants from tropical fog mountain","volume":"122","author":"Boanares","year":"2018","journal-title":"Plant Physiol. Bioch."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1002\/polb.20277","article-title":"Hydration mechanism of polysaccharides: A comparative study","volume":"43","author":"Despond","year":"2005","journal-title":"J. Polym. Sci. B Polym. Phys."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/S0144-8617(99)00113-7","article-title":"Effect of polymer chain end on sorption isotherm of water by chitosan","volume":"41","author":"Gocho","year":"2000","journal-title":"Carbohyd. Polym."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"11783","DOI":"10.1073\/pnas.1832434100","article-title":"Cell wall arabinan is essential for guard cell function","volume":"100","author":"Jones","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1263","DOI":"10.1002\/j.1537-2197.1981.tb07834.x","article-title":"The balance of forces generated by the water potential in the cell-wall-matrix: A model","volume":"68","author":"McClendon","year":"1981","journal-title":"Am. J. Bot."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1037","DOI":"10.4161\/psb.20768","article-title":"Probing the mechanical contributions of the pectin matrix","volume":"7","author":"Braybrook","year":"2012","journal-title":"Plant Signal. Behav."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1046\/j.1365-313x.2000.00742.x","article-title":"Probing expansin action using cellulose\/hemicellulose composites","volume":"22","author":"Whitney","year":"2000","journal-title":"Plant J."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"14664","DOI":"10.1073\/pnas.0605979103","article-title":"Crystal structure and activities of EXPB1 (Zea m 1), a \u03b2-expansin and group-1 pollen allergen from maize","volume":"103","author":"Yennawar","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1093\/aob\/mcm138","article-title":"Space and Time in the Plant Cell Wall: Relationships between Cell Types, Cell Wall Rheology and Cell Function","volume":"101","author":"Thompson","year":"2008","journal-title":"Ann. Bot."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1144","DOI":"10.1126\/science.205.4411.1144","article-title":"Determination of the pore size of cell walls of living plant cells","volume":"205","author":"Carpita","year":"1979","journal-title":"Science"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1291","DOI":"10.1093\/jexbot\/52.359.1291","article-title":"Extensiometric determination of the rheological properties of the epidermis of growing tomato fruit","volume":"52","author":"Thompson","year":"2001","journal-title":"J. Exp. Bot."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1711","DOI":"10.1093\/jxb\/45.Special_Issue.1711","article-title":"Autolysis and extension of isolated walls from growing cucumber hypocotyls","volume":"45","author":"Cosgrove","year":"1994","journal-title":"J. Exp. Bot."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Pereira, C.S., Ribeiro, J.M.L., Vatulescu, A.D., Findlay, K., MacDougall, A.J., and Jackson, P.A.P. (2011). Extensin network formation in Vitis vinifera callus cells is an essential and causal event in rapid and H2O2-induced reduction in primary cell wall hydration. BMC Plant Biol., 11.","DOI":"10.1186\/1471-2229-11-106"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"2085","DOI":"10.1073\/pnas.94.5.2085","article-title":"The MUR1 gene of Arabidopsis thaliana encodes an isoform of GDP-d-mannose-4,6-dehydratase, catalyzing the first step in the de novo synthesis\u2009of\u2009GDP-l-fucose","volume":"94","author":"Bonin","year":"1997","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"3340","DOI":"10.1073\/pnas.052450699","article-title":"The mur2 mutant of Arabidopsis thaliana lacks fucosylated xyloglucan because of a lesion in fucosyltransferase AtFUT1","volume":"99","author":"Vanzin","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"846","DOI":"10.1126\/science.1062319","article-title":"Requirement of borate cross-linking of cell wall rhamnogalacturonan II for Arabidopsis growth","volume":"294","author":"Eberhard","year":"2001","journal-title":"Science"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1146\/annurev.arplant.55.031903.141750","article-title":"Rhamnogalacturonan II: Structure and function of a borate cross-linked cell wall pectic polysaccharide","volume":"55","author":"Ishii","year":"2004","journal-title":"Ann. Rev. Plant Biol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1046\/j.1365-313X.1997.12020335.x","article-title":"Mutants of Arabidopsis thaliana with altered cell wall polysaccharide composition","volume":"12","author":"Reiter","year":"1997","journal-title":"Plant J."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1104\/pp.103.021873","article-title":"Tensile properties of Arabidopsis cell walls depend on both a xyloglucan cross-linked microfibrillar network and rhamnogalacturonan II-borate complexes","volume":"132","author":"Ryden","year":"2003","journal-title":"Plant Physiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1007\/s00425-004-1205-x","article-title":"L-Galactose replaces L-fucose in the pectic polysaccharide rhamnogalacturonan II synthesized by the L-fucose-deficient mur1 Arabidopsis mutant","volume":"219","author":"Reuhs","year":"2004","journal-title":"Planta"},{"key":"ref_31","first-page":"1425","article-title":"Two endogenous proteins that induce cell wall extension in Plants","volume":"4","author":"Durachko","year":"1992","journal-title":"Plant Cell"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1007\/s00425-004-1373-8","article-title":"Biophysical consequences of remodelling the neutral side chains of rhamnogalacturonan I in tubers of transgenic potatoes","volume":"220","author":"Ulvskov","year":"2005","journal-title":"Planta"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1016\/j.carbpol.2016.07.113","article-title":"Pectin impacts cellulose fibre architecture and hydrogel mechanics in the absence of calcium","volume":"153","author":"Bonilla","year":"2016","journal-title":"Carbohyd. Polym."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"10399","DOI":"10.1021\/ja504108h","article-title":"Water\u2013Polysaccharide Interactions in the Primary Cell Wall of Arabidopsis thaliana from Polarization Transfer Solid-State NMR","volume":"136","author":"White","year":"2014","journal-title":"J. Am. Chem. Soc."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"055005","DOI":"10.1088\/0965-0393\/23\/5\/055005","article-title":"Modeling the effects of pH and ionic strength on swelling of anionic polyelectrolyte gels","volume":"23","author":"Drozdov","year":"2015","journal-title":"Model. Simul. Mater. Sci. Eng."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/0008-6215(94)84228-0","article-title":"Selectivity and cooperativity in the binding of calcium ions by pectins","volume":"256","author":"Garnier","year":"1994","journal-title":"Carbohyd. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1016\/S0176-1617(99)80088-1","article-title":"Cell wall characteristics and structure of hydrated and dry leaves of the resurrection plant Craterostigma wilmsii, a microscopical study","volume":"155","author":"Sherwin","year":"1999","journal-title":"J. Plant Physiol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"739","DOI":"10.1007\/s00425-012-1785-9","article-title":"Arabinose-rich polymers as an evolutionary strategy to plasticize resurrection plant cell walls against desiccation","volume":"237","author":"Moore","year":"2013","journal-title":"Planta"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1168","DOI":"10.1016\/j.jplph.2007.09.006","article-title":"Water stress and cell wall polysaccharides in the apical root zone of wheat cultivars varying in drought tolerance","volume":"165","author":"Leucci","year":"2008","journal-title":"J. Plant Physiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/S0176-1617(97)80033-8","article-title":"Changes in amounts and molecular mass distribution of cell-wall polysaccharides of wheat (Triticum aestivum L.) coleoptiles under water stress","volume":"151","author":"Wakabayashi","year":"1997","journal-title":"J. Plant Physiol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1104\/pp.106.2.607","article-title":"Root growth maintenance at low water potentials (increased activity of xyloglucan endotransglycosylase and its possible regulation by abscisic acid)","volume":"106","author":"Wu","year":"1994","journal-title":"Plant Physiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1686","DOI":"10.3389\/fpls.2016.01686","article-title":"Drought and heat differentially affect XTH expression and XET activity and action in 3-day-old seedlings of durum wheat cultivars with different stress susceptibility","volume":"7","author":"Iurlaro","year":"2016","journal-title":"Front. Plant Sci."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"867","DOI":"10.1007\/s00299-010-0989-3","article-title":"Constitutive expression of CaXTH3, a hot pepper xyloglucan endotransglucosylase\/hydrolase, enhanced tolerance to salt and drought stresses without phenotypic defects in tomato plants (Solanum lycopersicum cv. Dotaerang)","volume":"30","author":"Choi","year":"2011","journal-title":"Plant Cell Rep."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Chen, Y., Zhang, B., Li, C., Lei, C., Kong, C., Yang, Y., and Gonget, M. (2019). A comprehensive expression analysis of the expansin gene family in potato (Solanum tuberosum) discloses stress-responsive expansin-like B genes for drought and heat tolerances. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0219837"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1104\/pp.111.3.765","article-title":"Growth maintenance of the maize primary root at low water potentials involves increases in cell wall extension properties, expansin activity and wall susceptibility to expansins","volume":"111","author":"Wu","year":"1996","journal-title":"Plant Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1471","DOI":"10.1104\/pp.126.4.1471","article-title":"Modification of expansin transcript levels in the maize primary root at low water potentials","volume":"126","author":"Wu","year":"2001","journal-title":"Plant Physiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1017\/S0960258520000379","article-title":"The expansin EXP1 gene in the elongation zone is induced during soybean embryonic axis germination and differentially expressed in response to ABA and PEG treatments","volume":"31","author":"Montechiarini","year":"2021","journal-title":"Seed Sci. Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"333","DOI":"10.2135\/cropsci2010.07.0391","article-title":"Cloning and characterization of a gene, AsEXP1, encoding expansin proteins inducible by heat stress and hormones in creeping bentgrass","volume":"51","author":"Zhou","year":"2011","journal-title":"Crop Sci."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1104\/pp.69.5.1145","article-title":"Complete turgor maintenance at low water potentials in the elongating region of maize leaves","volume":"69","author":"Michelena","year":"1982","journal-title":"Plant Physiol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1043","DOI":"10.1093\/jxb\/42.8.1043","article-title":"Turgor, growth and rheological gradients of wheat roots following osmotic stress","volume":"42","author":"Pritchard","year":"1991","journal-title":"J. Exp. Bot."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"438","DOI":"10.1104\/pp.96.2.438","article-title":"Spatial distribution of turgor and root growth at low water potentials","volume":"96","author":"Spollen","year":"1991","journal-title":"Plant Physiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1104\/pp.113.2.559","article-title":"Xylem Sap pH increase: A drought signal received at the apoplastic face of the guard cell that involves the suppression of saturable abscisic acid uptake by the epidermal symplast","volume":"113","author":"Wilkinson","year":"1997","journal-title":"Plant Physiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1016\/j.pbi.2008.03.006","article-title":"Pectin structure and biosynthesis","volume":"11","author":"Mohnen","year":"2008","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1698","DOI":"10.1104\/pp.126.4.1698","article-title":"Formation of rhamnogalacturonan II-borate dimer in pectin determines cell wall thickness of pumpkin tissue","volume":"126","author":"Ishii","year":"2001","journal-title":"Plant Physiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1401","DOI":"10.1104\/pp.117.4.1401","article-title":"The boron requirement and cell wall properties of growing and stationary suspension-cultured Chenopodium album L. cells","volume":"117","author":"Fleischer","year":"1998","journal-title":"Plant Physiol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1036","DOI":"10.1111\/tpj.14088","article-title":"Suppression of Arabidopsis GGLT1 affects growth by reducing the L-galactose content and borate cross-linking of rhamnogalacturonan-II","volume":"96","author":"Sechet","year":"2018","journal-title":"Plant J."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1111\/nph.15867","article-title":"Arabinose biosynthesis is critical for salt stress tolerance in Arabidopsis","volume":"224","author":"Zhao","year":"2019","journal-title":"New Phytol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.plantsci.2017.12.012","article-title":"Root cell wall solutions for crop plants in saline soils","volume":"269","author":"Byrt","year":"2018","journal-title":"Plant Sci."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1007\/s004250000319","article-title":"Expression of a heterologous expansin in transgenic tomato plants","volume":"211","author":"Rochange","year":"2000","journal-title":"Planta"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1016\/S0301-4622(02)00365-4","article-title":"Theoretical and experimental studies on freezing point depression and vapor pressure deficit as methods to measure osmotic pressure of aqueous polyethylene glycol and bovine serum albumin solutions","volume":"104","author":"Kiyosawa","year":"2003","journal-title":"Biophys. Chem."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1016\/j.bpc.2003.11.010","article-title":"Reappraisal of disparities between osmolality estimates by freezing point depression and vapour pressure deficit methods","volume":"107","author":"Winzor","year":"2004","journal-title":"Biophys. Chem."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/10\/7\/1263\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:20:37Z","timestamp":1760163637000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/10\/7\/1263"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,6,22]]},"references-count":61,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2021,7]]}},"alternative-id":["plants10071263"],"URL":"https:\/\/doi.org\/10.3390\/plants10071263","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,6,22]]}}}