fd0fc481849ff04d736a9146d560d8d9 jouexpbot_a2026m9v77n16p5067.pdf e234b54767b63f5317e228f0d4277b800f6cc16a jouexpbot_a2026m9v77n16p5067.pdf f8095edf207142c49fea45bea94845728d1f764fd1bc30dff04057f0ff4422fc jouexpbot_a2026m9v77n16p5067.pdf Title: Regulatory structural variations upstream of a NAC transcription factor control fruit maturity timing in peach Subject: DOI: 10.1093/jxb/erag210; Journal of Experimental Botany, 77, 16, 2026-04-29.; Abstract: Fruit maturity date is a key developmental trait in fruit trees, as it determines the timing of when fruits enter the ripening phase, thereby shaping the harvest calendar and fruit quality. In peach, ripening time varies over several months, yet the genetic and regulatory basis underlying this extensive phenological diversity has remained largely unresolved. Here, we dissect the long-standing chromosome 4 maturity date locus (qMD4.1) and show that a multi-allelic series of structural variants in the upstream regulatory region of the transcription factor NAC1 explains most of the observed variation in ripening time in cultivated peach. Across multiple independent populations and diverse genetic backgrounds, these variants (collectively termed Md) stratify maturity date from ultra-early to late cultivars with largely additive and dosage-dependent effects. Using recombinant-based genetic dissection, long-read sequencing, transcriptomic analyses, and epigenomic data integration, we show that Md alleles are consistently associated with allele-dependent shifts in the temporal dynamics of NAC1 transcript accumulation, and that the affected region overlaps with chromatin features characteristic of regulatory activity. By resolving the genetic architecture of a major phenological locus, our results support a central role for cis-regulatory structural variation in modulating developmental timing in peach and provide biologically interpretable markers for predicting maturity date. More broadly, this work illustrates how non-coding structural variation can contribute to variation in fruit developmental trajectories, with implications for breeding strategies aimed at modulating harvest windows in peach and other stone fruit species. Keywords: Fruit trees; maturity date; peach; QTL; ripening; structural variants Author: Alessandro Giulio Tagliabue 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: Thu Aug 20 17:50:28 2026 CEST ModDate: Mon Oct 19 11:28:20 2026 CEST Custom Metadata: yes Metadata Stream: yes Tagged: yes UserProperties: no Suspects: no Form: none JavaScript: no Pages: 16 Encrypted: no Page size: 612.283 x 790.866 pts Page rot: 0 File size: 1664328 bytes Optimized: no PDF version: 1.6 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- NWVXXB+MicrosoftSansSerif CID TrueType Identity-H yes yes yes 57 0 WXPZWT+HelveticaNeueLTStd-LtIt Type 1C Custom yes yes yes 43 0 KVNGDX+HelveticaNeueLTStd-Lt Type 1C Custom yes yes yes 42 0 RFETRN+HelveticaNeueLTStd-Md Type 1C Custom yes yes yes 1434 0 HAVICQ+HelveticaNeueLTStd-Bd Type 1C Custom yes yes yes 41 0 VDICDR+HelveticaNeueLTStd-BdIt Type 1C Custom yes yes yes 61 0 PJPRGK+ArialMT TrueType WinAnsi yes yes no 1 0 XXVIZT+BemboMTStd-Regular Type 1C Custom yes yes yes 44 0 VKFPFT+BemboMTStd-Italic Type 1C Custom yes yes yes 45 0 TTUORU+TimesNewRomanPSMT CID TrueType Identity-H yes yes yes 46 0 PLOFKM+STIXGeneral-Regular Type 1C Custom yes yes yes 47 0 KREPPL+TimesNewRomanPS-ItalicMT CID TrueType Identity-H yes yes yes 94 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2026-10-20 04:03:41 CEST RepresentationInformation: jouexpbot_a2026m9v77n16p5067.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-10-19 11:29:06 CEST Size: 1664328 Format: PDF Version: 1.6 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf Profile: Tagged PDF PDFMetadata: Objects: 1495 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: Regulatory structural variations upstream of a NAC transcription factor control fruit maturity timing in peach Destination: head1 Children: Item: Title: Introduction Destination: secerag210-s1 Item: Title: Materials and methods Destination: secerag210-s2 Children: Item: Title: Plant materials and phenotyping Destination: secerag210-s2.1 Item: Title: Genotyping, linkage map construction, and quantitative trait locus mapping Destination: secerag210-s2.2 Item: Title: DNA extraction and resequencing Destination: secerag210-s2.3 Item: Title: Genomics analyses Destination: secerag210-s2.4 Item: Title: Validation and genotyping of Md alleles Destination: secerag210-s2.5 Item: Title: Fruit sampling and RNA sequencing analyses Destination: secerag210-s2.6 Item: Title: Epigenomic datasets and data processing Destination: secerag210-s2.7 Item: Title: Results Destination: secerag210-s3 Children: Item: Title: Maturity date mapping in D×P progeny Destination: secerag210-s3.1 Item: Title: Genomic variation at the Md locus Destination: secerag210-s3.2 Item: Title: Association of Md alleles with maturity date across multiple segregating progenies Destination: secerag210-s3.3 Item: Title: Association of Md alleles with maturity date in germplasm collections Destination: secerag210-s3.4 Item: Title: Functional evidence of the effects of Md structural variants on NAC1 expression Destination: secerag210-s3.5 Item: Title: Origin and spread of Md alleles Destination: secerag210-s3.6 Item: Title: Discussion Destination: secerag210-s4 Children: Item: Title: Allelic determinants of maturity date in peach Destination: secerag210-s4.1 Item: Title: Functional evidence and transcriptional dynamics of the Md locus Destination: secerag210-s4.2 Item: Title: Breeding implications and marker-assisted selection Destination: secerag210-s4.3 Item: Title: Conclusion Destination: secerag210-s4.4 Item: Title: Supplementary data Destination: secerag210-s5 Item: Title: Acknowledgements Destination: acknow Item: Title: Author contributions Destination: secerag210-s6 Item: Title: Conflict of interest Destination: secerag210-s7 Item: Title: Funding Destination: secerag210-s8 Item: Title: Data availability Destination: secerag210-s9 Item: Title: References Destination: reflist Info: Title: Regulatory structural variations upstream of a NAC transcription factor control fruit maturity timing in peach Author: Alessandro Giulio Tagliabue Subject: DOI: 10.1093/jxb/erag210; Journal of Experimental Botany, 77, 16, 2026-04-29.; Abstract: Fruit maturity date is a key developmental trait in fruit trees, as it determines the timing of when fruits enter the ripening phase, thereby shaping the harvest calendar and fruit quality. In peach, ripening time varies over several months, yet the genetic and regulatory basis underlying this extensive phenological diversity has remained largely unresolved. Here, we dissect the long-standing chromosome 4 maturity date locus (qMD4.1) and show that a multi-allelic series of structural variants in the upstream regulatory region of the transcription factor NAC1 explains most of the observed variation in ripening time in cultivated peach. Across multiple independent populations and diverse genetic backgrounds, these variants (collectively termed Md) stratify maturity date from ultra-early to late cultivars with largely additive and dosage-dependent effects. Using recombinant-based genetic dissection, long-read sequencing, transcriptomic analyses, and epigenomic data integration, we show that Md alleles are consistently associated with allele-dependent shifts in the temporal dynamics of NAC1 transcript accumulation, and that the affected region overlaps with chromatin features characteristic of regulatory activity. By resolving the genetic architecture of a major phenological locus, our results support a central role for cis-regulatory structural variation in modulating developmental timing in peach and provide biologically interpretable markers for predicting maturity date. More broadly, this work illustrates how non-coding structural variation can contribute to variation in fruit developmental trajectories, with implications for breeding strategies aimed at modulating harvest windows in peach and other stone fruit species. Keywords: Fruit trees; maturity date; peach; QTL; ripening; structural variants 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: Thu Aug 20 18:20:28 CEST 2026 ModDate: Mon Oct 19 11:28:20 CEST 2026 ID: 0x3ceb091dc77b53502008368f74ed2c40, 0x894dd2be1953856c619785e94c759e76 Filters: FilterPipeline: FlateDecode FilterPipeline: DCTDecode Images: Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1162 ImageHeight: 754 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1129 ImageHeight: 448 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1162 ImageHeight: 1150 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Image: NisoImageMetadata: 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NWVXXB+MicrosoftSansSerif Flags: Nonsymbolic FontBBox: -580, -210, 1386, 922 FontFile2: true XMP: uuid:e44694d8-adb1-4146-9795-3a8169b7087c uuid:B3C760D1-D666-51F2-C062-C7F4B7654CD3 True Fruit trees; maturity date; peach; QTL; ripening; structural variants PDFlib+PDI 9.0.7p3 (C++/Win64); modified using iTextSharp.LGPLv2.Core 3.7.4.0 application/pdf 10.1093/jxb/erag210 en Oxford University Press Regulatory structural variations upstream of a NAC transcription factor control fruit maturity timing in peach Alessandro Giulio Tagliabue James Friel Remo Chiozzotto Irina Baccichet Cassia da Silva Linge Elisa Calastri Gianluca Biffi Nathalia Zaracho Stefano Gattolin Sabrina Micali Iban Eduardo Daniele Bassi Laura Rossini Marco Cirilli DOI: 10.1093/jxb/erag210; Journal of Experimental Botany, 77, 16, 2026-04-29.; Abstract: Fruit maturity date is a key developmental trait in fruit trees, as it determines the timing of when fruits enter the ripening phase, thereby shaping the harvest calendar and fruit quality. In peach, ripening time varies over several months, yet the genetic and regulatory basis underlying this extensive phenological diversity has remained largely unresolved. Here, we dissect the long-standing chromosome 4 maturity date locus (qMD4.1) and show that a multi-allelic series of structural variants in the upstream regulatory region of the transcription factor NAC1 explains most of the observed variation in ripening time in cultivated peach. Across multiple independent populations and diverse genetic backgrounds, these variants (collectively termed Md) stratify maturity date from ultra-early to late cultivars with largely additive and dosage-dependent effects. Using recombinant-based genetic dissection, long-read sequencing, transcriptomic analyses, and epigenomic data integration, we show that Md alleles are consistently associated with allele-dependent shifts in the temporal dynamics of NAC1 transcript accumulation, and that the affected region overlaps with chromatin features characteristic of regulatory activity. By resolving the genetic architecture of a major phenological locus, our results support a central role for cis-regulatory structural variation in modulating developmental timing in peach and provide biologically interpretable markers for predicting maturity date. More broadly, this work illustrates how non-coding structural variation can contribute to variation in fruit developmental trajectories, with implications for breeding strategies aimed at modulating harvest windows in peach and other stone fruit species. Fruit trees maturity date peach QTL ripening structural variants © The Author(s) 2026. Published by Oxford University Press on behalf of the Society for Experimental Biology. Journal Journal of Experimental Botany © The Author(s) 2026. 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