0fbb1c737551cbd32007498e3a7c7e89 plaphy_a2025m10v199n2pkiaf420.pdf 6d466fb82b7246a5f7b3616ed440e406f8df26d7 plaphy_a2025m10v199n2pkiaf420.pdf 532843c303b83c08bb99d28ff3368ff82e7d1f43776332a77f301565b7bba1af plaphy_a2025m10v199n2pkiaf420.pdf Title: Glycosylated sterols enhance cold tolerance in tomato via membrane stabilization and jasmonate signaling Subject: DOI: 10.1093/plphys/kiaf420; Plant Physiology, 199, 2, 2025-09-23.; Abstract: Free and glycosylated sterols play a central role in maintaining the structural integrity and proper functioning of the plasma membrane, which serves as the primary sensor of cold and initiates signaling cascades to mitigate chilling-induced damage. Here, we characterize the cold-response of tomato (Solanum lycopersicum cv. Micro-Tom) mutants with higher and lower ratios of glycosylated to free sterols than wild-type plants, resulting from the overexpression and silencing of the STEROL GLYCOSYLTRANSFERASES 1 and 2 (SlSGT2 and SlSGT1), respectively. The SlSGT2 overexpressing mutants show increased cold tolerance, membrane stability, and oxidative stress responses, while silencing of the SlSGT1 gene causes the opposite phenotypes. Furthermore, changes in the glycosylated to free sterols ratio activate distinct transcriptional programs that establish a preconditioned stress-responsive state under basal conditions and trigger a more efficient response to cold in the SlSGT2 overexpressing mutants, as well as compromise the capacity to withstand the effects of cold stress in the SlSGT1-silenced mutants. The SlSGT2 overexpressing mutants also contain higher levels of jasmonates under basal conditions and display enhanced biosynthesis of these hormones under cold stress compared to SlSGT1-silenced and control plants. The facilitating effect of elevated glycosylated to free sterols ratio on jasmonic acid (JA) biosynthesis and signaling leads to the activation of cold-responsive genes, including those of the CBF–COR pathway, antioxidant defenses, and polyamine biosynthesis. Our findings provide key insights into the mechanisms by which glycosylated sterols help improve cold tolerance in tomato. Keywords: Author: Cuiyun Deng Creator: Servigistics Arbortext Advanced Print Publisher 11.1.4546/W-x64 Producer: PDFlib+PDI 9.0.7p3 (C++/Win64); modified using iTextSharp.LGPLv2.Core 3.7.4.0 CreationDate: Fri Oct 10 08:33:57 2025 CEST ModDate: Wed Sep 16 15:56:51 2026 CEST Custom Metadata: yes Metadata Stream: yes Tagged: yes UserProperties: no Suspects: no Form: none JavaScript: no Pages: 15 Encrypted: no Page size: 595.276 x 782.362 pts Page rot: 0 File size: 3549723 bytes Optimized: no PDF version: 1.6 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- POIXQZ+TimesNewRomanPSMT CID TrueType Identity-H yes yes yes 98 0 EWGJMM+CaeciliaLTStd-LightItalic Type 1C Custom yes yes yes 39 0 FNCZJA+CaeciliaLTStd-Light Type 1C Custom yes yes yes 37 0 COSLZQ+CaeciliaLTStd-Bold Type 1C Custom yes yes yes 36 0 XTFEVQ+CaeciliaLTStd-Heavy Type 1C Custom yes yes yes 38 0 FGVSLO+ArialMT TrueType WinAnsi yes yes no 1 0 COUHEC+STIXGeneral-Regular Type 1C Custom yes yes yes 40 0 Helvetica-Bold Type 1 Custom no no no 100 0 Helvetica-Bold Type 1 Custom no no no 110 0 Helvetica-Bold Type 1 Custom no no no 129 0 Helvetica-Bold Type 1 Custom no no no 144 0 Helvetica-Bold Type 1 Custom no no no 160 0 Helvetica-Bold Type 1 Custom no no no 193 0 Helvetica-Bold Type 1 Custom no no no 252 0 DBFRCZ+TimesNewRomanPS-ItalicMT CID TrueType Identity-H yes yes yes 286 0 IEKZHD+CaeciliaLTStd-BoldItalic Type 1C Custom yes yes yes 287 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2026-09-17 04:14:01 CEST RepresentationInformation: plaphy_a2025m10v199n2pkiaf420.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-09-16 15:57:54 CEST Size: 3549723 Format: PDF Version: 1.6 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf Profile: Tagged PDF PDFMetadata: Objects: 1489 FreeObjects: 1 IncrementalUpdates: 0 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: true NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: OneColumn PageMode: UseOutlines Language: en Outlines: Item: Title: Glycosylated sterols enhance cold tolerance in tomato via membrane stabilization and jasmonate signaling Destination: head1 Children: Item: Title: Introduction Destination: seckiaf420-s1 Item: Title: Results Destination: seckiaf420-s2 Children: Item: Title: Differential impact of contrasting membrane sterol profiles on tomato cold tolerance Destination: seckiaf420-s2.1 Item: Title: The imbalance of membrane sterols alters the response to cold-induced oxidative damage Destination: seckiaf420-s2.2 Item: Title: Transcriptome-wide analysis of SGT1ami and SGT2ox plants uncovers antagonistic acclimation states to cold treatment Destination: seckiaf420-s2.3 Item: Title: Membrane sterol imbalance alters polyamines and jasmonates levels Destination: seckiaf420-s2.4 Item: Title: Discussion Destination: seckiaf420-s3 Item: Title: Materials and methods Destination: seckiaf420-s4 Children: Item: Title: Plant growth conditions and cold treatment Destination: seckiaf420-s4.1 Children: Item: Title: Generation of transgenic lines overexpressing SlSGT2 Destination: seckiaf420-s4.1.1 Item: Title: RNA isolation and quantitative real-time PCR analysis Destination: seckiaf420-s4.2 Item: Title: Sterol analysis Destination: seckiaf420-s4.3 Item: Title: Electrolyte leakage assay Destination: seckiaf420-s4.4 Item: Title: Chlorophyll content determination Destination: seckiaf420-s4.5 Item: Title: Maximum quantum efficiency of PSII (Fv/Fm) Destination: seckiaf420-s4.6 Item: Title: Histochemical detection of reactive oxygen Species (ROS) Destination: seckiaf420-s4.7 Item: Title: MDA quantification Destination: seckiaf420-s4.8 Item: Title: Enzyme activity assays Destination: seckiaf420-s4.9 Item: Title: Transcriptome profiling and analysis Destination: seckiaf420-s4.10 Item: Title: Quantification of jasmonates Destination: seckiaf420-s4.11 Item: Title: Polyamine analysis Destination: seckiaf420-s4.12 Item: Title: Statistical analysis Destination: seckiaf420-s4.13 Item: Title: Accession numbers Destination: seckiaf420-s4.14 Item: Title: Acknowledgments Destination: acknow Item: Title: Author contributions Destination: seckiaf420-s5 Item: Title: Supplementary data Destination: seckiaf420-s6 Item: Title: Funding Destination: seckiaf420-s7 Item: Title: References Destination: reflist Info: Title: Glycosylated sterols enhance cold tolerance in tomato via membrane stabilization and jasmonate signaling Author: Cuiyun Deng Subject: DOI: 10.1093/plphys/kiaf420; Plant Physiology, 199, 2, 2025-09-23.; Abstract: Free and glycosylated sterols play a central role in maintaining the structural integrity and proper functioning of the plasma membrane, which serves as the primary sensor of cold and initiates signaling cascades to mitigate chilling-induced damage. Here, we characterize the cold-response of tomato (Solanum lycopersicum cv. Micro-Tom) mutants with higher and lower ratios of glycosylated to free sterols than wild-type plants, resulting from the overexpression and silencing of the STEROL GLYCOSYLTRANSFERASES 1 and 2 (SlSGT2 and SlSGT1), respectively. The SlSGT2 overexpressing mutants show increased cold tolerance, membrane stability, and oxidative stress responses, while silencing of the SlSGT1 gene causes the opposite phenotypes. Furthermore, changes in the glycosylated to free sterols ratio activate distinct transcriptional programs that establish a preconditioned stress-responsive state under basal conditions and trigger a more efficient response to cold in the SlSGT2 overexpressing mutants, as well as compromise the capacity to withstand the effects of cold stress in the SlSGT1-silenced mutants. The SlSGT2 overexpressing mutants also contain higher levels of jasmonates under basal conditions and display enhanced biosynthesis of these hormones under cold stress compared to SlSGT1-silenced and control plants. The facilitating effect of elevated glycosylated to free sterols ratio on jasmonic acid (JA) biosynthesis and signaling leads to the activation of cold-responsive genes, including those of the CBF COR pathway, antioxidant defenses, and polyamine biosynthesis. Our findings provide key insights into the mechanisms by which glycosylated sterols help improve cold tolerance in tomato. Keywords: Creator: Servigistics Arbortext Advanced Print Publisher 11.1.4546/W-x64 Producer: PDFlib+PDI 9.0.7p3 (C++/Win64); modified using iTextSharp.LGPLv2.Core 3.7.4.0 CreationDate: Fri Oct 10 09:03:57 CEST 2025 ModDate: Wed Sep 16 15:56:51 CEST 2026 ID: 0x32883b8e3ba8d0a598626adef2660c37, 0xe339147497103f8b67aab1b78ced2681 Filters: FilterPipeline: FlateDecode FilterPipeline: DCTDecode Images: Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 666 ImageHeight: 751 BitsPerSample: 8 BitsPerSampleUnit: integer Fonts: Type0: Font: BaseFont: POIXQZ+TimesNewRomanPSMT Encoding: Identity-H ToUnicode: true Font: BaseFont: DBFRCZ+TimesNewRomanPS-ItalicMT Encoding: Identity-H ToUnicode: true Type1: Font: BaseFont: COSLZQ+CaeciliaLTStd-Bold FontSubset: true FirstChar: 1 LastChar: 62 FontDescriptor: FontName: COSLZQ+CaeciliaLTStd-Bold Flags: Nonsymbolic, ForceBold FontBBox: -166, -273, 1141, 938 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: FNCZJA+CaeciliaLTStd-Light FontSubset: true FirstChar: 1 LastChar: 95 FontDescriptor: FontName: FNCZJA+CaeciliaLTStd-Light Flags: Nonsymbolic FontBBox: -164, -266, 1130, 908 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: XTFEVQ+CaeciliaLTStd-Heavy FontSubset: true FirstChar: 1 LastChar: 30 FontDescriptor: FontName: XTFEVQ+CaeciliaLTStd-Heavy Flags: Nonsymbolic, ForceBold FontBBox: -170, -277, 1146, 970 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: EWGJMM+CaeciliaLTStd-LightItalic FontSubset: true FirstChar: 1 LastChar: 65 FontDescriptor: FontName: EWGJMM+CaeciliaLTStd-LightItalic Flags: Nonsymbolic, Italic FontBBox: -166, -267, 1033, 909 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: COUHEC+STIXGeneral-Regular FontSubset: true FirstChar: 1 LastChar: 10 FontDescriptor: FontName: COUHEC+STIXGeneral-Regular Flags: Nonsymbolic FontBBox: -995, -455, 2000, 1055 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: IEKZHD+CaeciliaLTStd-BoldItalic FontSubset: true FirstChar: 1 LastChar: 5 FontDescriptor: FontName: IEKZHD+CaeciliaLTStd-BoldItalic Flags: Nonsymbolic, Italic, ForceBold FontBBox: -169, -273, 1082, 932 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true TrueType: Font: BaseFont: FGVSLO+ArialMT FontSubset: true FirstChar: 32 LastChar: 121 FontDescriptor: FontName: FGVSLO+ArialMT Flags: Nonsymbolic FontBBox: -664, -324, 2000, 1039 FontFile2: true Encoding: WinAnsiEncoding CIDFontType2: Font: BaseFont: DBFRCZ+TimesNewRomanPS-ItalicMT CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: DBFRCZ+TimesNewRomanPS-ItalicMT Flags: Nonsymbolic, Italic FontBBox: -498, -307, 1333, 1023 FontFile2: true Font: BaseFont: POIXQZ+TimesNewRomanPSMT CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: POIXQZ+TimesNewRomanPSMT Flags: Nonsymbolic FontBBox: -568, -307, 2046, 1040 FontFile2: true XMP: uuid:d7a1beba-7a19-4272-af5e-522adc58190e uuid:D0DCE567-E528-9503-0A29-CC52890E62E1 True PDFlib+PDI 9.0.7p3 (C++/Win64); modified using iTextSharp.LGPLv2.Core 3.7.4.0 application/pdf 10.1093/plphys/kiaf420 en Oxford University Press Glycosylated sterols enhance cold tolerance in tomato via membrane stabilization and jasmonate signaling Cuiyun Deng Antoni Garcia-Molina Aurelio Gómez-Cadenas Vicente Vives-Peris Rubén Alcázar Albert Ferrer Teresa Altabella DOI: 10.1093/plphys/kiaf420; Plant Physiology, 199, 2, 2025-09-23.; Abstract: Free and glycosylated sterols play a central role in maintaining the structural integrity and proper functioning of the plasma membrane, which serves as the primary sensor of cold and initiates signaling cascades to mitigate chilling-induced damage. Here, we characterize the cold-response of tomato (Solanum lycopersicum cv. Micro-Tom) mutants with higher and lower ratios of glycosylated to free sterols than wild-type plants, resulting from the overexpression and silencing of the STEROL GLYCOSYLTRANSFERASES 1 and 2 (SlSGT2 and SlSGT1), respectively. The SlSGT2 overexpressing mutants show increased cold tolerance, membrane stability, and oxidative stress responses, while silencing of the SlSGT1 gene causes the opposite phenotypes. Furthermore, changes in the glycosylated to free sterols ratio activate distinct transcriptional programs that establish a preconditioned stress-responsive state under basal conditions and trigger a more efficient response to cold in the SlSGT2 overexpressing mutants, as well as compromise the capacity to withstand the effects of cold stress in the SlSGT1-silenced mutants. The SlSGT2 overexpressing mutants also contain higher levels of jasmonates under basal conditions and display enhanced biosynthesis of these hormones under cold stress compared to SlSGT1-silenced and control plants. The facilitating effect of elevated glycosylated to free sterols ratio on jasmonic acid (JA) biosynthesis and signaling leads to the activation of cold-responsive genes, including those of the CBF–COR pathway, antioxidant defenses, and polyamine biosynthesis. Our findings provide key insights into the mechanisms by which glycosylated sterols help improve cold tolerance in tomato. © The Author(s) 2025. Published by Oxford University Press on behalf of American Society of Plant Biologists. Journal Plant Physiology © The Author(s) 2025. 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