{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T19:13:57Z","timestamp":1792782837525,"version":"4.5.0"},"reference-count":101,"publisher":"Springer Science and Business Media LLC","license":[{"start":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T00:00:00Z","timestamp":1789776000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T00:00:00Z","timestamp":1789776000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"University of Innsbruck and Medical University of Innsbruck","id":[{"id":"https:\/\/ror.org\/054pv6659","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Protoplasma"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Cytorrhysis describes the shrinkage of plant cells caused by water loss, during which the protoplast and cell wall contract while remaining connected. Cytorrhysis can occur naturally during drought and freezing and can be experimentally induced by exposure to osmotic substances. Although traditionally studied separately, the cytorrhysis process itself shares the same underlying mechanisms: water efflux along chemical potential gradients and the mechanical response of the cell wall\u2013plasma membrane continuum. Cytorrhysis has been documented across diverse taxa \u2013 including algae, bryophytes, ferns and seed plants \u2013 indicating that it represents a fundamental response of walled cells to dehydration. Besides osmotic effects, the extent of cytorrhysis is also strongly affected by biomechanical cell structure. Cell size, wall thickness, elasticity and tissue architecture influence the likelihood of cellular collapse. Small, thick-walled cells withstand larger pressure differences and develop negative turgor pressure whereas large, thin-walled cells collapse more readily. During freezing, extracellular ice imposes temperature-dependent dehydration forces, and biomechanical constraints may generate negative turgor pressure. Recent advances, including cryo-microscopy, differential scanning calorimetry and psychrometric water-potential measurements, allow more precise quantification of freeze dehydration and structural responses. Functionally, cytorrhysis can be protective or damaging. Moderate, reversible cytorrhysis helps maintain membrane integrity e.g. in desiccation-tolerant species. In freezing environments, cells undergoing cytorrhysis survive temperatures far below those tolerated by supercooling cells. However, excessive dehydration may cause cell death. Integrating classical observations with modern biophysical approaches provides a renewed perspective on cytorrhysis and highlights structural traits that could be exploited to enhance plant resilience to drought and freezing stress in a changing climate.<\/jats:p>","DOI":"10.1007\/s00709-026-02263-1","type":"journal-article","created":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T01:25:43Z","timestamp":1789781143000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Cytorrhysis under drought, osmotic and freezing stress"],"prefix":"10.1007","author":[{"given":"Matthias","family":"Stegner","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karin","family":"Fehringer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexander","family":"Fl\u00f6rl","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna-Lena","family":"Strasser","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jasmin","family":"Lindner","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sandra","family":"Plangger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tanja","family":"Sch\u00e4fernolte","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Viktoria","family":"Thoma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gilbert","family":"Neuner","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ingeborg","family":"Lang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,9,19]]},"reference":[{"key":"2263_CR1","first-page":"619","volume":"165","author":"W Abel","year":"1956","unstructured":"Abel W (1956) Die Austrocknungsresistenz der Laubmoose. Sitzungsberichte der Akademie der Wissenschaften mathematisch-naturwissenschaftliche Klasse 165:619\u2013707","journal-title":"Sitzungsberichte der Akademie der Wissenschaften mathematisch-naturwissenschaftliche Klasse"},{"key":"2263_CR2","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1016\/j.plantsci.2018.03.002","volume":"270","author":"R Arora","year":"2018","unstructured":"Arora R (2018) Mechanism of freeze-thaw injury and recovery: A cool retrospective and warming up to new ideas. Plant Sci 270:301\u2013313. https:\/\/doi.org\/10.1016\/j.plantsci.2018.03.002","journal-title":"Plant Sci"},{"key":"2263_CR3","first-page":"83","volume":"10","author":"E Asahina","year":"1956","unstructured":"Asahina E (1956) The freezing process of plant cells. Contrib Inst Low Temp Sci 10:83\u2013126","journal-title":"Contrib Inst Low Temp Sci"},{"key":"2263_CR4","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1104\/pp.76.1.201","volume":"76","author":"EN Ashworth","year":"1984","unstructured":"Ashworth EN, Abeles FB (1984) Freezing behavior of water in small pores and the possible role in the freezing of plant tissues. Plant Physiol 76:201\u2013204. https:\/\/doi.org\/10.1104\/pp.76.1.201","journal-title":"Plant Physiol"},{"issue":"3","key":"2263_CR5","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1007\/s12038-007-0049-5","volume":"32","author":"EH Beck","year":"2007","unstructured":"Beck EH, Fettig S, Knake C, Hartig K, Bhattarai T (2007) Specific and unspecific responses of plants to cold and drought stress. J Biosci 32(3):501\u2013510. https:\/\/doi.org\/10.1007\/s12038-007-0049-5","journal-title":"J Biosci"},{"issue":"2","key":"2263_CR6","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1006\/anbo.1996.0318","volume":"79","author":"RP Beckett","year":"1997","unstructured":"Beckett RP (1997) Pressure\u2013volume analysis of a range of poikilohydric plants implies the existence of negative turgor in vegetative cells. Ann Botany 79(2):145\u2013152. https:\/\/doi.org\/10.1006\/anbo.1996.0318","journal-title":"Ann Bot"},{"issue":"3","key":"2263_CR7","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1046\/j.1365-313X.1991.t01-11-00999.x","volume":"1","author":"G Bianchi","year":"1991","unstructured":"Bianchi G, Gamba A, Murelli C, Salamini F, Bartels D (1991) Novel carbohydrate metabolism in the resurrection plant Craterostigma plantagineum. Plant J 1(3):355\u2013359. https:\/\/doi.org\/10.1046\/j.1365-313X.1991.t01-11-00999.x","journal-title":"Plant J"},{"issue":"4","key":"2263_CR8","doi-asserted-by":"publisher","first-page":"300","DOI":"10.1007\/BF01514018","volume":"89","author":"R Biebl","year":"1940","unstructured":"Biebl R (1940) Einige zellphysiologische Beobachtungen an Hookeria lucens (L.) Sm. Oesterr Bot Z 89(4):300\u2013306","journal-title":"Oesterr Bot Z"},{"issue":"1\u20134","key":"2263_CR9","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1007\/s00709-009-0053-8","volume":"243","author":"O Buchner","year":"2010","unstructured":"Buchner O, Neuner G (2010) Freezing cytorrhysis and critical temperature thresholds for photosystem II in the peat moss Sphagnum capillifolium. Protoplasma 243(1\u20134):63\u201371. https:\/\/doi.org\/10.1007\/s00709-009-0053-8","journal-title":"Protoplasma"},{"issue":"1","key":"2263_CR10","doi-asserted-by":"publisher","DOI":"10.1186\/s13007-020-00586-5","volume":"16","author":"O Buchner","year":"2020","unstructured":"Buchner O, Steiner P, Andosch A, Holzinger A, Stegner M, Neuner G, Lutz-Meindl U (2020) A new technical approach for preparing frozen biological samples for electron microscopy. Plant Methods 16(1):16. https:\/\/doi.org\/10.1186\/s13007-020-00586-5","journal-title":"Plant Methods"},{"issue":"4411","key":"2263_CR11","doi-asserted-by":"publisher","first-page":"1144","DOI":"10.1126\/science.205.4411.1144","volume":"205","author":"N Carpita","year":"1979","unstructured":"Carpita N, Sabularse D, Montezinos D, Delmer DP (1979) Determination of the pore size of cell walls of living plant cells. Science 205(4411):1144\u20131147. https:\/\/doi.org\/10.1126\/science.205.4411.1144","journal-title":"Science"},{"issue":"15","key":"2263_CR12","doi-asserted-by":"publisher","first-page":"4075","DOI":"10.1093\/jxb\/erx243","volume":"68","author":"X Cheng","year":"2017","unstructured":"Cheng X, Lang I, Adeniji OS, Griffing L (2017) Plasmolysis-deplasmolysis causes changes in endoplasmic reticulum form, movement, flow, and cytoskeletal association. J Exp Bot 68(15):4075\u20134087. https:\/\/doi.org\/10.1093\/jxb\/erx243","journal-title":"J Exp Bot"},{"issue":"2","key":"2263_CR13","doi-asserted-by":"publisher","first-page":"378","DOI":"10.1111\/nph.12829","volume":"203","author":"Y Ding","year":"2014","unstructured":"Ding Y, Zhang Y, Zheng QS, Tyree MT (2014) Pressure-volume curves: revisiting the impact of negative turgor during cell collapse by literature review and simulations of cell micromechanics. New Phytol 203(2):378\u2013387. https:\/\/doi.org\/10.1111\/nph.12829","journal-title":"New Phytol"},{"issue":"4","key":"2263_CR14","doi-asserted-by":"publisher","first-page":"1139","DOI":"10.1104\/pp.75.4.1139","volume":"75","author":"MF Dowgert","year":"1984","unstructured":"Dowgert MF, Steponkus PL (1984) Behavior of the plasma membrane of isolated protoplasts during a freeze-thaw cycle. Plant Physiol 75(4):1139\u20131151. https:\/\/doi.org\/10.1104\/pp.75.4.1139","journal-title":"Plant Physiol"},{"issue":"1","key":"2263_CR15","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1046\/j.1365-2818.1997.1360696.x","volume":"185","author":"WE England","year":"1997","unstructured":"England WE, McCully ME, Huang CX (1997) Solvent vapour lock: an extreme case of the problems caused by lignified and suberized cell walls during resin infiltration. J Microsc 185(1):85\u201393. https:\/\/doi.org\/10.1046\/j.1365-2818.1997.1360696.x","journal-title":"J Microsc"},{"issue":"2","key":"2263_CR16","doi-asserted-by":"publisher","first-page":"212","DOI":"10.3390\/plants9020212","volume":"9","author":"I Ezquer","year":"2020","unstructured":"Ezquer I, Salameh I, Colombo L, Kalaitzis P (2020) Plant cell walls tackling climate change: Insights into plant cell wall remodeling, its regulation, and biotechnological strategies to improve crop adaptations and photosynthesis in response to global warming. Plants 9(2):212. https:\/\/doi.org\/10.3390\/plants9020212","journal-title":"Plants"},{"issue":"6","key":"2263_CR17","doi-asserted-by":"publisher","first-page":"291","DOI":"10.1007\/BF01502879","volume":"8","author":"HW Fischer","year":"1911","unstructured":"Fischer HW (1911) Das Wasser im Plasma. Z f\u00fcr Chemie und Ind der Kolloide 8(6):291\u2013302. https:\/\/doi.org\/10.1007\/BF01502879","journal-title":"Z f\u00fcr Chemie und Ind der Kolloide"},{"issue":"11","key":"2263_CR18","doi-asserted-by":"publisher","first-page":"2049","DOI":"10.1007\/s00018-019-03388-8","volume":"77","author":"N Gigli-Bisceglia","year":"2020","unstructured":"Gigli-Bisceglia N, Engelsdorf T, Hamann T (2020) Plant cell wall integrity maintenance in model plants and crop species-relevant cell wall components and underlying guiding principles. Cell Mol Life Sci 77(11):2049\u20132077. https:\/\/doi.org\/10.1007\/s00018-019-03388-8","journal-title":"Cell Mol Life Sci"},{"issue":"5","key":"2263_CR19","doi-asserted-by":"publisher","first-page":"707","DOI":"10.1104\/pp.56.5.707","volume":"56","author":"LV Gusta","year":"1975","unstructured":"Gusta LV, Burke MJ, Kapoor AC (1975) Determination of unfrozen water in winter cereals at subfreezing temperatures. Plant Physiol 56(5):707\u2013709. https:\/\/doi.org\/10.1104\/pp.56.5.707","journal-title":"Plant Physiol"},{"issue":"4","key":"2263_CR20","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1007\/s004420100628","volume":"126","author":"UG Hacke","year":"2001","unstructured":"Hacke UG, Sperry JS, Pockman WT, Davis SD, McCulloh KA (2001) Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure. Oecologia 126(4):457\u2013461. https:\/\/doi.org\/10.1007\/s004420100628","journal-title":"Oecologia"},{"issue":"1","key":"2263_CR21","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1111\/j.1438-8677.1988.tb00014.x","volume":"101","author":"J Hansen","year":"1988","unstructured":"Hansen J, Beck E (1988) Evidence for ideal and non-ideal equilibrium freezing of leaf water in frosthardy Ivy (Hedera helix) and Winter Barley (Hordeum vulgare). Bot Acta 101(1):76\u201382. https:\/\/doi.org\/10.1111\/j.1438-8677.1988.tb00014.x","journal-title":"Bot Acta"},{"key":"2263_CR22","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1007\/978-3-642-80446-5_11","volume-title":"Progress in Botany","author":"W Hartung","year":"1998","unstructured":"Hartung W, Schiller P, Dietz K (1998) Physiology of poikilohydric plants. In: Behnke HD, Esser K, Kadereit JW, L\u00fcttge U, Runge M (eds) Progress in Botany, vol vol 59. Springer, Berlin, Heidelberg, pp 299\u2013327. https:\/\/doi.org\/10.1007\/978-3-642-80446-5_11"},{"key":"2263_CR23","doi-asserted-by":"crossref","unstructured":"H\u00f6fler K (1920) Ein Schema f\u00fcr die osmotische Leistung der Pflanzenzelle. Ber. d. d. Bot. Ges., 38","DOI":"10.1111\/j.1438-8677.1920.tb07879.x"},{"key":"2263_CR24","unstructured":"H\u00f6fler K (1947) Einige Nekrosen bei F\u00e4rbung mit Akridinorange. Sitzungsberichte der Akademie der Wissenschaften mathematisch-naturwissenschaftliche Klasse (156):585\u2013644"},{"issue":"2","key":"2263_CR25","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1093\/oxfordjournals.aob.a086688","volume":"53","author":"RM Hoggart","year":"1984","unstructured":"Hoggart RM, Clarke AE (1984) Porosity of gladiolus stigmatic papillae and pollen tube walls. Ann Botany 53(2):271\u2013278. https:\/\/doi.org\/10.1093\/oxfordjournals.aob.a086688","journal-title":"Ann Bot"},{"issue":"1\u20133","key":"2263_CR26","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1016\/s0367-1615(17)31719-6","volume":"108","author":"H Holle","year":"1915","unstructured":"Holle H (1915) Untersuchungen \u00fcber Welken, Vertrocknen und Wieder-straffwerden. Flora oder Allgemeine Bot Ztg 108(1\u20133):73\u2013126. https:\/\/doi.org\/10.1016\/s0367-1615(17)31719-6","journal-title":"Flora oder Allgemeine Bot Ztg"},{"issue":"1","key":"2263_CR27","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1007\/BF02674817","volume":"20","author":"W Iljin","year":"1933","unstructured":"Iljin W (1933) \u00dcber den K\u00e4ltetod der Pflanzen und seine Ursachen. Protoplasma 20(1):105\u2013124","journal-title":"Protoplasma"},{"issue":"1","key":"2263_CR28","doi-asserted-by":"publisher","first-page":"546","DOI":"10.1104\/pp.104.058404","volume":"139","author":"AL Jacobsen","year":"2005","unstructured":"Jacobsen AL, Ewers FW, Pratt RB, Paddock WA, Davis SD (2005) Do xylem fibers affect vessel cavitation resistance? Plant Physiol 139(1):546\u2013556. https:\/\/doi.org\/10.1104\/pp.104.058404","journal-title":"Plant Physiol"},{"issue":"8","key":"2263_CR29","doi-asserted-by":"publisher","first-page":"1172","DOI":"10.1038\/s41477-024-01742-8","volume":"10","author":"Y Jaillais","year":"2024","unstructured":"Jaillais Y, Bayer E, Bergmann DC, Botella MA, Boutt\u00e9 Y, Bozkurt TO, Caillaud M-C, Germain V, Grossmann G, Heilmann I, Hemsley PA, Kirchhelle C, Martini\u00e8re A, Miao Y, Mongrand S, M\u00fcller S, Noack LC, Oda Y, Ott T, Pan X, Pleskot R, Potocky M, Robert S, Rodriguez CS, Simon-Plas F, Russinova E, Van Damme D, Van Norman JM, Weijers D, Yalovsky S, Yang Z, Zelazny E, Gronnier J (2024) Guidelines for naming and studying plasma membrane domains in plants. Nat Plants 10(8):1172\u20131183. https:\/\/doi.org\/10.1038\/s41477-024-01742-8","journal-title":"Nat Plants"},{"issue":"12","key":"2263_CR30","doi-asserted-by":"publisher","first-page":"3101","DOI":"10.1111\/pce.13078","volume":"40","author":"E Kuprian","year":"2017","unstructured":"Kuprian E, Munkler C, Resnyak A, Zimmermann S, Tuong TD, Gierlinger N, M\u00fcller T, Livingston D, Neuner G (2017) Complex bud architecture and cell-specific chemical patterns enable supercooling of Picea abies bud primordia. Plant Cell Environ 40(12):3101\u20133112. https:\/\/doi.org\/10.1111\/pce.13078","journal-title":"Plant Cell Environ"},{"issue":"2","key":"2263_CR31","doi-asserted-by":"publisher","first-page":"736","DOI":"10.1104\/pp.011601","volume":"131","author":"K Kuroda","year":"2003","unstructured":"Kuroda K, Kasuga J, Arakawa K, Fujikawa S (2003) Xylem ray parenchyma cells in boreal hardwood species respond to subfreezing temperatures by deep supercooling that is accompanied by incomplete desiccation. Plant Physiol 131(2):736\u2013744. https:\/\/doi.org\/10.1104\/pp.011601","journal-title":"Plant Physiol"},{"key":"2263_CR32","volume-title":"Grundpraktikum zur Pflanzenphysiologie: Online Version 2019","author":"U Kutschera","year":"2019","unstructured":"Kutschera U (2019) Grundpraktikum zur Pflanzenphysiologie: Online Version 2019. Quelle & Meyer, Wiesbaden"},{"issue":"1","key":"2263_CR33","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/S0044-328X(77)80129-3","volume":"82","author":"E Kyriakopoulos","year":"1977","unstructured":"Kyriakopoulos E, Richter H (1977) A comparison of methods for the determination of water status in Quercus ilex L. Z Pflanzenphysiol 82(1):14\u201327. https:\/\/doi.org\/10.1016\/S0044-328X(77)80129-3","journal-title":"Z Pflanzenphysiol"},{"issue":"4","key":"2263_CR34","doi-asserted-by":"publisher","first-page":"583","DOI":"10.3390\/plants3040583","volume":"3","author":"I Lang","year":"2014","unstructured":"Lang I, Sassmann S, Schmidt B, Komis G (2014) Plasmolysis: Loss of turgor and beyond. Plants 3(4):583\u2013593. https:\/\/doi.org\/10.3390\/plants3040583","journal-title":"Plants"},{"key":"2263_CR35","volume-title":"\u00d6kophysiologie der Pflanzen","author":"W Larcher","year":"1994","unstructured":"Larcher W (1994) \u00d6kophysiologie der Pflanzen. Ulmer, Stuttgart"},{"issue":"2","key":"2263_CR36","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1016\/S0367-1615(17)31891-8","volume":"132","author":"E Lau\u00e9","year":"1938","unstructured":"Lau\u00e9 E (1938) Untersuchungen an Pflanzenzellen im Dampfraum. Flora 132(2):193\u2013224. https:\/\/doi.org\/10.1016\/S0367-1615(17)31891-8","journal-title":"Flora"},{"issue":"10","key":"2263_CR37","doi-asserted-by":"publisher","first-page":"1731","DOI":"10.1111\/j.1365-3040.2010.02178.x","volume":"33","author":"T Lenn\u00e9","year":"2010","unstructured":"Lenn\u00e9 T, Bryant G, Hocart CH, Huang CX, Ball MC (2010) Freeze avoidance: a dehydrating moss gathers no ice. Plant Cell Environ 33(10):1731\u20131741. https:\/\/doi.org\/10.1111\/j.1365-3040.2010.02178.x","journal-title":"Plant Cell Environ"},{"key":"2263_CR38","volume-title":"Responses of plants to environmental stresses. Chilling, freezing, and high temperature stresses","author":"J Levitt","year":"1980","unstructured":"Levitt J (1980) Responses of plants to environmental stresses. Chilling, freezing, and high temperature stresses, vol 1. Academic Press , London, New York"},{"issue":"3","key":"2263_CR39","doi-asserted-by":"publisher","first-page":"1207","DOI":"10.1104\/pp.107.111351","volume":"146","author":"D Li\u00e9nard","year":"2008","unstructured":"Li\u00e9nard D, Durambur G, Kiefer-Meyer M-C, Nogu\u00e9 F, Menu-Bouaouiche L, Charlot F, Gomord V, Lassalles J-P (2008) Water transport by aquaporins in the extant plant Physcomitrella patens. Plant Physiol 146(3):1207\u20131218. https:\/\/doi.org\/10.1104\/pp.107.111351","journal-title":"Plant Physiol"},{"issue":"11","key":"2263_CR40","doi-asserted-by":"publisher","first-page":"3807","DOI":"10.1093\/jxb\/erac108","volume":"73","author":"J Liu","year":"2022","unstructured":"Liu J, Willick IR, Hiraki H, Forand AD, Lawrence JR, Swerhone GDW, Wei YD, Ghosh S, Lee YK, Olsen JE, Usadel B, Wormit A, Gunl M, Karunakaran C, Dynes JJ, Tanino KK (2022) Cold and exogenous calcium alter Allium fistulosum cell wall pectin to depress intracellular freezing temperatures. J Exp Bot 73(11):3807\u20133822. https:\/\/doi.org\/10.1093\/jxb\/erac108","journal-title":"J Exp Bot"},{"issue":"1","key":"2263_CR41","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1016\/j.tca.2007.04.019","volume":"459","author":"D Livingston","year":"2007","unstructured":"Livingston D (2007) Quantifying liquid water in frozen plant tissues by isothermal calorimetry. Thermochimica acta 459(1):116\u2013120. https:\/\/doi.org\/10.1016\/j.tca.2007.04.019","journal-title":"Thermochimica acta"},{"key":"2263_CR42","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-662-67530-4","volume-title":"Experimentelle Pflanzen\u00f6kologie","author":"R Matyssek","year":"2020","unstructured":"Matyssek R, Herppich WB (2020) Experimentelle Pflanzen\u00f6kologie. Springer. https:\/\/doi.org\/10.1007\/978-3-662-67530-4"},{"issue":"5","key":"2263_CR43","doi-asserted-by":"publisher","first-page":"901","DOI":"10.1093\/aob\/mcl133","volume":"98","author":"JD Mauseth","year":"2006","unstructured":"Mauseth JD (2006) Structure\u2013function relationships in highly modified shoots of Cactaceae. Ann Botany 98(5):901\u2013926. https:\/\/doi.org\/10.1093\/aob\/mcl133","journal-title":"Ann Bot"},{"issue":"2","key":"2263_CR44","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1085\/jgp.47.2.347","volume":"47","author":"P Mazur","year":"1963","unstructured":"Mazur P (1963) Kinetics of water loss from cells at subzero temperatures and likelihood of intracellular freezing. J Gen Physiol 47(2):347\u2013369. https:\/\/doi.org\/10.1085\/jgp.47.2.347","journal-title":"J Gen Physiol"},{"issue":"2","key":"2263_CR45","doi-asserted-by":"publisher","first-page":"658","DOI":"10.1111\/j.1749-6632.1965.tb45420.x","volume":"125","author":"P Mazur","year":"1965","unstructured":"Mazur P (1965) The role of cell membranes in the freezing of yeast and other single cells. Ann N Y Acad Sci 125(2):658\u2013676. https:\/\/doi.org\/10.1111\/j.1749-6632.1965.tb45420.x","journal-title":"Ann N Y Acad Sci"},{"issue":"5","key":"2263_CR46","doi-asserted-by":"publisher","first-page":"665","DOI":"10.1093\/aob\/mch191","volume":"94","author":"ME McCully","year":"2004","unstructured":"McCully ME, Canny MJ, Huang CX (2004) The management of extracellular ice by petioles of frost-resistant herbaceous plants. Ann Botany 94(5):665\u2013674. https:\/\/doi.org\/10.1093\/aob\/mch191","journal-title":"Ann Botany"},{"issue":"4","key":"2263_CR47","doi-asserted-by":"publisher","first-page":"119","DOI":"10.2307\/1546753","volume":"70","author":"JH Miller","year":"1980","unstructured":"Miller JH (1980) Differences in the apparent permeability of spore walls and prothallial cell walls in Onoclea sensibilis. Am Fern J 70(4):119\u2013123. https:\/\/doi.org\/10.2307\/1546753","journal-title":"Am Fern J"},{"key":"2263_CR48","volume-title":"Untersuchungen \u00fcber das Erfrieren der Pflanzen","author":"H Molisch","year":"1897","unstructured":"Molisch H (1897) Untersuchungen \u00fcber das Erfrieren der Pflanzen. G. Fischer, Jena"},{"issue":"3","key":"2263_CR49","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1007\/s00425-012-1785-9","volume":"237","author":"JP Moore","year":"2013","unstructured":"Moore JP, Nguema-Ona EE, Vicr\u00e9-Gibouin M, S\u00f8rensen I, Willats WGT, Driouich A, Farrant JM (2013) Arabinose-rich polymers as an evolutionary strategy to plasticize resurrection plant cell walls against desiccation. Planta 237(3):739\u2013754. https:\/\/doi.org\/10.1007\/s00425-012-1785-9","journal-title":"Planta"},{"key":"2263_CR50","first-page":"435","volume":"15","author":"H M\u00fcller-Thurgau","year":"1886","unstructured":"M\u00fcller-Thurgau H (1886) \u00dcber das Gefrieren und Erfrieren der Pflanzen. Landw Jahrbucher 15:435\u2013610","journal-title":"Landw Jahrbucher"},{"issue":"12","key":"2263_CR51","doi-asserted-by":"publisher","first-page":"1539","DOI":"10.1046\/j.1365-3040.1999.00508.x","volume":"22","author":"G Neuner","year":"1999","unstructured":"Neuner G, Braun V, Buchner O, Taschler D (1999) Leaf rosette closure in the alpine rock species Saxifraga paniculata Mill.: significance for survival of drought and heat under high irradiation. Plant Cell Environ 22(12):1539\u20131548. https:\/\/doi.org\/10.1046\/j.1365-3040.1999.00508.x","journal-title":"Plant Cell Environ"},{"issue":"4","key":"2263_CR52","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1016\/S0176-1617(85)80022-5","volume":"121","author":"JJ Oertli","year":"1985","unstructured":"Oertli JJ (1985) The response of plant cells to different forms of moisture stress. J Plant Physiol 121(4):295\u2013300. https:\/\/doi.org\/10.1016\/S0176-1617(85)80022-5","journal-title":"J Plant Physiol"},{"issue":"3\u20134","key":"2263_CR53","doi-asserted-by":"publisher","first-page":"365","DOI":"10.1016\/S0176-1617(86)80048-7","volume":"124","author":"JJ Oertli","year":"1986","unstructured":"Oertli JJ (1986) The effect of cell size on cell collapse under negative turgor pressure. J Plant Physiol 124(3\u20134):365\u2013370. https:\/\/doi.org\/10.1016\/S0176-1617(86)80048-7","journal-title":"J Plant Physiol"},{"key":"2263_CR54","first-page":"73","volume-title":"Structural and functional responses to environmental stresses: water shortage","author":"JJ Oertli","year":"1989","unstructured":"Oertli JJ (1989) The plant cell\u2019s resistance to consequences of negative turgor pressure. In: Kreeb KH (ed) Structural and functional responses to environmental stresses: water shortage. SPB Academic Publishing, The Hague, pp 73\u201388"},{"key":"2263_CR55","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1017\/CBO9780511753305.004","volume-title":"Water transport in plants under climatic stress","author":"JJ Oertli","year":"1993","unstructured":"Oertli JJ (1993) Effect of cavitation on the status of water in plants. In: Borgetti M, Grace J, Raschi I (eds) Water transport in plants under climatic stress. Cambridge University Press, Cambridge, pp 27\u201340. https:\/\/doi.org\/10.1017\/CBO9780511753305.004"},{"issue":"2","key":"2263_CR56","first-page":"281","volume":"11","author":"J Oertli","year":"1990","unstructured":"Oertli J, Lips S, Agami M (1990) The strength of sclerophyllous cells to resist collapse due to negative turgor pressure. Acta Oecol 11(2):281\u2013289","journal-title":"Acta Oecol"},{"issue":"1","key":"2263_CR57","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1146\/annurev.pp.18.060167.002131","volume":"18","author":"CR Olien","year":"1967","unstructured":"Olien CR (1967) Freezing stresses and survival. Annu Rev Plant Physiol 18(1):387\u2013408. https:\/\/doi.org\/10.1146\/annurev.pp.18.060167.002131","journal-title":"Annu Rev Plant Physiol"},{"issue":"4","key":"2263_CR58","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1111\/j.1469-8137.1994.tb02952.x","volume":"126","author":"KJ Oparka","year":"1994","unstructured":"Oparka KJ (1994) Plasmolysis - New insights into an old process. New Phytol 126(4):571\u2013591. https:\/\/doi.org\/10.1111\/j.1469-8137.1994.tb02952.x","journal-title":"New Phytol"},{"issue":"4","key":"2263_CR59","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1002\/9781119312994.apr0746","volume":"3","author":"PE Panter","year":"2020","unstructured":"Panter PE, Panter JR, Knight H (2020) Impact of cell-wall structure and composition on plant freezing tolerance. Annual Plant Reviews Online 3(4):607\u2013642. https:\/\/doi.org\/10.1002\/9781119312994.apr0746","journal-title":"Annual Plant Reviews Online"},{"issue":"4","key":"2263_CR60","doi-asserted-by":"publisher","first-page":"1933","DOI":"10.1104\/pp.111.192880","volume":"158","author":"YB Park","year":"2012","unstructured":"Park YB, Cosgrove DJ (2012) A revised architecture of primary cell walls based on biomechanical changes induced by substrate-specific endoglucanases. Plant Physiol 158(4):1933\u20131943. https:\/\/doi.org\/10.1104\/pp.111.192880","journal-title":"Plant Physiol"},{"issue":"2","key":"2263_CR61","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1086\/335761","volume":"114","author":"J Parker","year":"1952","unstructured":"Parker J (1952) Desiccation in conifer leaves: Anatomical changes and determination of the lethal level. Bot Gaz 114(2):189\u2013198. https:\/\/doi.org\/10.1086\/335761","journal-title":"Bot Gaz"},{"issue":"4","key":"2263_CR62","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1006\/anbo.2000.1352","volume":"87","author":"RS Pearce","year":"2001","unstructured":"Pearce RS (2001) Plant freezing and damage. Ann Botany 87(4):417\u2013424. https:\/\/doi.org\/10.1006\/anbo.2000.1352","journal-title":"Ann Botany"},{"key":"2263_CR63","first-page":"664","volume":"84","author":"M Pfeiffer","year":"1933","unstructured":"Pfeiffer M (1933) Frostuntersuchungen an Fichtentrieben. Tharandter forstliche Jahrbuch 84:664\u2013695","journal-title":"Tharandter forstliche Jahrbuch"},{"issue":"328","key":"2263_CR64","doi-asserted-by":"publisher","first-page":"1845","DOI":"10.1093\/jxb\/49.328.1845","volume":"49","author":"MCF Proctor","year":"1998","unstructured":"Proctor MCF, Nagy Z, Csintalan Z, Tak\u00e1cs Z (1998) Water-content components in bryophytes: analysis of pressure-volume relationships. J Exp Bot 49(328):1845\u20131854. https:\/\/doi.org\/10.1093\/jxb\/49.328.1845","journal-title":"J Exp Bot"},{"issue":"2","key":"2263_CR65","doi-asserted-by":"publisher","first-page":"597","DOI":"10.1104\/pp.111.2.597","volume":"111","author":"C Rajashekar","year":"1996","unstructured":"Rajashekar C, Burke MJ (1996) Freezing characteristics of rigid plant tissues - development of cell tension during extracellular freezing. Plant Physiol 111(2):597\u2013603. https:\/\/doi.org\/10.1104\/pp.111.2.597","journal-title":"Plant Physiol"},{"issue":"2","key":"2263_CR66","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1104\/pp.111.2.605","volume":"111","author":"CB Rajashekar","year":"1996","unstructured":"Rajashekar CB, Lafta A (1996) Cell-wall changes and cell tension in response to cold acclimation and exogenous abscisic acid in leaves and cell cultures. Plant Physiol 111(2):605\u2013612. https:\/\/doi.org\/10.1104\/pp.111.2.605","journal-title":"Plant Physiol"},{"key":"2263_CR67","doi-asserted-by":"publisher","unstructured":"Ralser M, Stegner M, Neuner G (2024) When water turns to ice: Control of ice volume and freezing dynamics as important aspects of cold acclimation. Environ Exp Bot 227. https:\/\/doi.org\/10.1016\/j.envexpbot.2024.105957","DOI":"10.1016\/j.envexpbot.2024.105957"},{"key":"2263_CR68","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1007\/BF01911870","volume":"18","author":"O Renner","year":"1932","unstructured":"Renner O (1932) Zur Kenntnis des Wasserhaushalts javanischer Kleinepiphyten. Planta 18:215\u2013287. https:\/\/doi.org\/10.1007\/BF01911870","journal-title":"Planta"},{"issue":"3","key":"2263_CR69","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1111\/j.1399-3054.1997.tb00566.x","volume":"99","author":"S Rhizopoulou","year":"1997","unstructured":"Rhizopoulou S (1997) Is negative turgor fallacious? Physiol Plant 99(3):505\u2013510. https:\/\/doi.org\/10.1111\/j.1399-3054.1997.tb00566.x","journal-title":"Physiol Plant"},{"key":"2263_CR70","unstructured":"Sachs J (1860) Krystallbildung bei dem Gefrieren und Ver\u00e4nderung der Zellh\u00e4ute bei dem Aufthauen saftiger Pflanzenteile. Berichte \u00fcber die Verhandlungen der K\u00f6niglich-S\u00e4chsischen Gesellschaft der Wissenschaften zu Leipzig, Mathematisch-Physische Klasse, vol 12. K\u00f6niglich S\u00e4chsische Gesellschaft der Wissenschaften"},{"key":"2263_CR71","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-642-71745-1","volume-title":"Frost survival of plants","year":"1987","unstructured":"Sakai A, Larcher W (1987) Frost survival of plants. Ecological studies, vol 62. Springer, Berlin, Heidelberg. https:\/\/doi.org\/10.1007\/978-3-642-71745-1"},{"issue":"1","key":"2263_CR72","doi-asserted-by":"publisher","first-page":"190","DOI":"10.1104\/pp.83.1.190","volume":"83","author":"JE Schmidt","year":"1987","unstructured":"Schmidt JE, Kaiser WM (1987) Response of the succulent leaves of Peperomia magnoliaefolia to dehydration: Water relations and solute movement in chlorenchyma and hydrenchyma. Plant Physiol 83(1):190\u2013194. https:\/\/doi.org\/10.1104\/pp.83.1.190","journal-title":"Plant Physiol"},{"key":"2263_CR73","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-662-49880-4","volume-title":"Pflanzenphysiologie","author":"P Schopfer","year":"2010","unstructured":"Schopfer P, Brennicke A (2010) Pflanzenphysiologie, vol 7. Spektrum Akademischer Verlag, Heidelberg. https:\/\/doi.org\/10.1007\/978-3-662-49880-4"},{"key":"2263_CR74","doi-asserted-by":"publisher","first-page":"7042","DOI":"10.3390\/ijms21197042","volume":"21","author":"M Stegner","year":"2020","unstructured":"Stegner M, Lackner B, Sch\u00e4fernolte T, Buchner O, Xiao N, Gierlinger N, Holzinger A, Neuner G (2020a) Winter nights during summer time: Stress physiological response to ice and facilitation of freezing cytorrhysis by elastic cell wall components in leaves of a nival species. Int J Mol Sci 21:7042. https:\/\/doi.org\/10.3390\/ijms21197042","journal-title":"Int J Mol Sci"},{"issue":"1","key":"2263_CR75","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13007-020-00617-1","volume":"16","author":"M Stegner","year":"2020","unstructured":"Stegner M, Wagner J, Neuner G (2020b) Ice accommodation in plant tissues pinpointed by cryo-microscopy in reflected-polarised-light. Plant Methods 16(1):1\u20139. https:\/\/doi.org\/10.1186\/s13007-020-00617-1","journal-title":"Plant Methods"},{"issue":"6","key":"2263_CR76","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.13793","volume":"174","author":"M Stegner","year":"2022","unstructured":"Stegner M, Fl\u00f6rl A, Lindner J, Plangger S, Schaefernolte T, Strasser AL, Thoma V, Walde J, Neuner G (2022) Freeze dehydration vs. supercooling of mesophyll cells: impact of cell wall, cellular and tissue traits on the extent of water displacement. Physiol Plant 174(6):e13793. https:\/\/doi.org\/10.1111\/ppl.13793","journal-title":"Physiol Plant"},{"key":"2263_CR77","doi-asserted-by":"publisher","DOI":"10.1016\/j.envexpbot.2024.105895","author":"M Stegner","year":"2024","unstructured":"Stegner M, Strasser AL, Neuner G (2024) Supercooling cells of frost hardy palm leaves: quantified percentage of frozen water and displacement from thermodynamic equilibrium. Environ Exp Bot. https:\/\/doi.org\/10.1016\/j.envexpbot.2024.105895","journal-title":"Environ Exp Bot"},{"key":"2263_CR78","doi-asserted-by":"publisher","unstructured":"Stegner M, Ralser M, Charalambous P, Tiloca G, Gierlinger N, Neuner G, Livingston D (2025) Differences in freezing dynamics in the tip and base of wheat (Triticum aestivum L.) leaves result in a difference in cold hardiness. Plant Stress 100853. https:\/\/doi.org\/10.1016\/j.stress.2025.100853","DOI":"10.1016\/j.stress.2025.100853"},{"key":"2263_CR79","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1111\/j.1438-8677.1900.tb04927.x","volume":"18","author":"C Steinbrinck","year":"1900","unstructured":"Steinbrinck C (1900a) \u00dcber die Grenzen des Schrumpfelns. Ber Dtsch Bot Ges 18:386\u2013396","journal-title":"Ber Dtsch Bot Ges"},{"key":"2263_CR80","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1111\/j.1438-8677.1900.tb04899.x","volume":"18","author":"C Steinbrinck","year":"1900","unstructured":"Steinbrinck C (1900b) Zur Terminologie der Volum\u00e4nderungen pflanzlicher Gewebe und organischer Substanzen bei wechselndem Fl\u00fcssigkeitsgehalt. Ber Dtsch Bot Ges 18:217\u2013224","journal-title":"Ber Dtsch Bot Ges"},{"issue":"2","key":"2263_CR81","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1111\/j.1438-8677.1910.tb06851.x","volume":"28","author":"C Steinbrinck","year":"1910","unstructured":"Steinbrinck C (1910) Weiteres \u00fcber den Koh\u00e4sionsmechanismus von Laubmoosbl\u00e4ttern. Ber Dtsch Bot Ges 28(2):19\u201330","journal-title":"Ber Dtsch Bot Ges"},{"key":"2263_CR82","doi-asserted-by":"publisher","unstructured":"Steiner P, Obwegeser S, Wanner G, Buchner O, L\u00fctz-Meindl U, Holzinger A (2020) Cell wall reinforcements accompany chilling and freezing stress in the streptophyte green alga Klebsormidium crenulatum. Front Plant Sci 11. https:\/\/doi.org\/10.3389\/fpls.2020.00873","DOI":"10.3389\/fpls.2020.00873"},{"issue":"17","key":"2263_CR83","doi-asserted-by":"publisher","first-page":"5203","DOI":"10.1093\/jxb\/erw279","volume":"67","author":"D Takahashi","year":"2016","unstructured":"Takahashi D, Kawamura Y, Uemura M (2016) Cold acclimation is accompanied by complex responses of glycosylphosphatidylinositol (GPI)-anchored proteins in Arabidopsis. J Exp Bot 67(17):5203\u20135215. https:\/\/doi.org\/10.1093\/jxb\/erw279","journal-title":"J Exp Bot"},{"issue":"12","key":"2263_CR84","doi-asserted-by":"publisher","first-page":"1858","DOI":"10.1093\/pcp\/pcab103","volume":"62","author":"D Takahashi","year":"2021","unstructured":"Takahashi D, Willick I, Kasuga J, Livingston D (2021) Responses of the plant cell wall to sub-zero temperatures: a brief update. Plant Cell Physiol 62(12):1858\u20131866. https:\/\/doi.org\/10.1093\/pcp\/pcab103","journal-title":"Plant Cell Physiol"},{"issue":"6","key":"2263_CR85","doi-asserted-by":"publisher","first-page":"1681","DOI":"10.1007\/s00709-019-01404-z","volume":"256","author":"K Trumh\u00f3va","year":"2019","unstructured":"Trumh\u00f3va K, Holzinger A, Obwegeser S, Neuner G, Pichrtova M (2019) The conjugating green alga Zygnema sp. (Zygnematophyceae) from the Arctic shows high frost tolerance in mature cells (pre-akinetes). Protoplasma 256(6):1681\u20131694. https:\/\/doi.org\/10.1007\/s00709-019-01404-z","journal-title":"Protoplasma"},{"issue":"23","key":"2263_CR86","doi-asserted-by":"publisher","first-page":"2738","DOI":"10.1139\/b76-294","volume":"54","author":"M Tyree","year":"1976","unstructured":"Tyree M (1976) Negative turgor pressure in plant cells: fact or fallacy? Can J Bot 54(23):2738\u20132746. https:\/\/doi.org\/10.1139\/b76-294","journal-title":"Can J Bot"},{"key":"2263_CR87","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1007\/978-3-642-68150-9_3","volume-title":"Physiological plant ecology II: water relations and carbon assimilation","author":"M Tyree","year":"1982","unstructured":"Tyree M, Jarvis P (1982) Water in tissues and cells. Physiological plant ecology II: water relations and carbon assimilation. Springer, pp 35\u201377"},{"issue":"1","key":"2263_CR88","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1104\/pp.109.1.15","volume":"109","author":"M Uemura","year":"1995","unstructured":"Uemura M, Joseph R, Steponkus PL (1995) Cold acclimation of Arabidopsis thaliana (Effect on plasma membrane lipid composition and freeze-induced lesions). Plant Physiol 109(1):15\u201330. https:\/\/doi.org\/10.1104\/pp.109.1.15","journal-title":"Plant Physiol"},{"key":"2263_CR89","doi-asserted-by":"publisher","DOI":"10.3203\/IWF\/E-1912","volume-title":"Osmotische Erscheinungen bei Pflanzenzellen: Plasmolyse, Zytorrhyse, Hookeria lucens (Musci)","author":"Url WG","year":"1973","unstructured":"Url WG (1973) Osmotische Erscheinungen bei Pflanzenzellen: Plasmolyse, Zytorrhyse, Hookeria lucens (Musci). Encyclopedia Cinematographica. IWF, G\u00f6ttingen, Germany. https:\/\/doi.org\/10.3203\/IWF\/E-1912"},{"key":"2263_CR90","doi-asserted-by":"publisher","unstructured":"Url WG (1974) Plasmolyse und Zytorrhyse. G\u00f6ttingen, Germany. https:\/\/doi.org\/10.3203\/iwf\/c-1144","DOI":"10.3203\/iwf\/c-1144"},{"key":"2263_CR91","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1111\/j.1365-313X.2005.02593.x","volume":"45","author":"PE Verslues","year":"2006","unstructured":"Verslues PE, Agarwal M, Katiyar-Agarwal S, Zhu J, Zhu J-K (2006) Methods and concepts in quantifying resistance to drought, salt and freezing, abiotic stresses that affect plant water status. Plant J 45:523\u2013539. https:\/\/doi.org\/10.1111\/j.1365-313X.2005.02593.x","journal-title":"Plant J"},{"issue":"11","key":"2263_CR92","doi-asserted-by":"publisher","first-page":"1329","DOI":"10.1111\/j.1365-3040.2004.01212.x","volume":"27","author":"M Vicr\u00e9","year":"2004","unstructured":"Vicr\u00e9 M, Farrant JM, Driouich A (2004) Insights into the cellular mechanisms of desiccation tolerance among angiosperm resurrection plant species. Plant Cell Environ 27(11):1329\u20131340. https:\/\/doi.org\/10.1111\/j.1365-3040.2004.01212.x","journal-title":"Plant Cell Environ"},{"issue":"41","key":"2263_CR93","doi-asserted-by":"publisher","first-page":"16444","DOI":"10.1073\/pnas.1316290110","volume":"110","author":"T Wang","year":"2013","unstructured":"Wang T, Park YB, Caporini MA, Rosay M, Zhong L, Cosgrove DJ, Hong M (2013) Sensitivity-enhanced solid-state NMR detection of expansin\u2019s target in plant cell walls. Proc Natl Acad Sci 110(41):16444\u201316449. https:\/\/doi.org\/10.1073\/pnas.1316290110","journal-title":"Proc Natl Acad Sci"},{"key":"2263_CR94","first-page":"732","volume":"26","author":"LS Weiss","year":"1991","unstructured":"Weiss LS, Palta JP (1991) Influence of thaw rates on the level of injury by freeze-thaw stress in a cold hardy potato species. HortScience 26:732\u2013733","journal-title":"HortScience"},{"key":"2263_CR95","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1104\/pp.62.5.699","volume":"62","author":"SC Wiest","year":"1978","unstructured":"Wiest SC, Steponkus PL (1978) Freeze-thaw injury to isolated spinach protoplasts and its simulation at above freezing temperatures. Plant Physiol 62:699\u2013705. https:\/\/doi.org\/10.1104\/pp.62.5.699","journal-title":"Plant Physiol"},{"issue":"22","key":"2263_CR96","doi-asserted-by":"publisher","first-page":"19404","DOI":"10.1074\/jbc.M011242200","volume":"276","author":"WG Willats","year":"2001","unstructured":"Willats WG, Orfila C, Limberg G, Buchholt HC, van Alebeek GJ, Voragen AG, Marcus SE, Christensen TM, Mikkelsen JD, Murray BS, Knox JP (2001) Modulation of the degree and pattern of methyl-esterification of pectic homogalacturonan in plant cell walls. Implications for pectin methyl esterase action, matrix properties, and cell adhesion. J Biol Chem 276(22):19404\u201319413. https:\/\/doi.org\/10.1074\/jbc.M011242200","journal-title":"J Biol Chem"},{"issue":"5","key":"2263_CR97","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1007\/BF00202015","volume":"9","author":"M Wisniewski","year":"1995","unstructured":"Wisniewski M, Davis G (1995) Immunogold localization of pectins and glycoproteins in tissues of peach with reference to deep supercooling. Trees 9(5):253\u2013260. https:\/\/doi.org\/10.1007\/BF00202015","journal-title":"Trees"},{"issue":"5","key":"2263_CR98","doi-asserted-by":"publisher","first-page":"770","DOI":"10.1007\/s00425-002-0814-5","volume":"215","author":"T Yamada","year":"2002","unstructured":"Yamada T, Kuroda K, Jitsuyama Y, Takezawa D, Arakawa K, Fujikawa S (2002) Roles of the plasma membrane and the cell wall in the responses of plant cells to freezing. Planta 215(5):770\u2013778. https:\/\/doi.org\/10.1007\/s00425-002-0814-5","journal-title":"Planta"},{"issue":"3","key":"2263_CR99","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1111\/pce.12860","volume":"40","author":"D Yang","year":"2017","unstructured":"Yang D, Li J, Ding Y, Tyree MT (2017a) Experimental evidence for negative turgor pressure in small leaf cells of Robinia pseudoacacia L. versus large cells of Metasequoia glyptostroboides Hu et W.C. Cheng. 2. H\u00f6fler diagrams below the volume of zero turgor and the theoretical implication for pressure-volume curves of living cells. Plant Cell Environ 40(3):340\u2013350. https:\/\/doi.org\/10.1111\/pce.12860","journal-title":"Plant Cell Environ"},{"issue":"3","key":"2263_CR100","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1111\/pce.12861","volume":"40","author":"DM Yang","year":"2017","unstructured":"Yang DM, Pan SA, Ding YT, Tyree MT (2017b) Experimental evidence for negative turgor pressure in small leaf cells of Robinia pseudoacacia L. versus large cells of Metasequoia glyptostroboides Hu et W.C.Cheng. Evidence from pressure-volume curve analysis of dead tissue. Plant Cell Environ 40(3):351\u2013363. https:\/\/doi.org\/10.1111\/pce.12861","journal-title":"Plant Cell Environ"},{"key":"2263_CR101","doi-asserted-by":"publisher","first-page":"1146","DOI":"10.1104\/pp.97.3.1146","volume":"97","author":"JJ Zhu","year":"1991","unstructured":"Zhu JJ, Beck E (1991) Water relations of Pachysandra leaves during freezing and thawing. Plant Physiol 97:1146\u20131153. https:\/\/doi.org\/10.1104\/pp.97.3.1146","journal-title":"Plant Physiol"}],"container-title":["Protoplasma"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00709-026-02263-1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00709-026-02263-1","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00709-026-02263-1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T01:25:46Z","timestamp":1789781146000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00709-026-02263-1"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,19]]},"references-count":101,"alternative-id":["2263"],"URL":"https:\/\/doi.org\/10.1007\/s00709-026-02263-1","relation":{},"ISSN":["0033-183X","1615-6102"],"issn-type":[{"value":"0033-183X","type":"print"},{"value":"1615-6102","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,19]]},"assertion":[{"value":"31 March 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 September 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"19 September 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}]}}