# Dynamic chromatin regulatory programs of sucrose and citric acid metabolism during fruit ripening in Citrus

**Authors:** Xin Song, Ting-Ting Wang, Peng Zhao, Li-Gang He, Yan-Jie Fan, Yu Zhang, Ting Liu, Chun-Mei Shi, Ying-Chun Jiang, Feng-Quan Tan, Abdelhafid Bendahmane, Chun-Long Li, Li-Ming Wu, Fang Song

PMC · DOI: 10.1093/plcell/koag060 · 2026-03-25

## TL;DR

This study explores how citrus fruits ripen by examining changes in chromatin and gene activity, revealing key regulators of sugar and acid metabolism.

## Contribution

The study identifies a regulatory cascade involving ABA, CrbZIP5b, and CrHSFA6B that controls citrus fruit ripening.

## Key findings

- A transcriptional regulatory network involving 52 key transcription factors was constructed for citrus ripening.
- CrHSFA6B and CrbZIP5b regulate genes involved in sucrose and citric acid metabolism during ripening.
- Abscisic acid (ABA) plays a central role in modulating citrus fruit ripening processes.

## Abstract

Fruit ripening is a complex, tightly regulated process that affects fruit flavor, nutritional value, and shelf life. Citrus is a typical non-climacteric fruit, as citrus fruits ripen independently of ethylene. However, the precise regulatory mechanism underlying this process remains to be elucidated. To dissect the epigenetic and transcriptional basis of citrus ripening, we performed integrative analyses of genome-wide histone modifications (H3K27ac, H3K4me3, and H3K27me3), chromatin accessibility, and transcriptome profiles throughout fruit ripening in Citrus reticulata. We constructed a transcriptional regulatory network encompassing 52 key transcription factors that orchestrate citrus fruit ripening. Further, we identified the key role of abscisic acid (ABA) in modulating sucrose and citric acid metabolism and uncovered the underlying transcriptional and epigenetic regulatory network during fruit ripening in C. reticulata. The findings demonstrate that CrHSFA6B and CrbZIP5b interact with the promoters of CrACO1 (Aconitase 1) and CrACO3 and CrSPSF1 (Sucrose Phosphate Synthase F1), regulating their expression to affect the sucrose and citric acid metabolism and fruit ripening. This study provides insights into the epigenomic dynamics of citrus fruit ripening and identifies a putative regulatory cascade centered on ABA, CrbZIP5b, and CrHSFA6B that modulates key processes in citrus ripening.

A transcriptional regulatory network of the citrus ripening process is established based on the histone modifications, chromatin accessibility, and transcriptome profiles. Two transcription factors activate sucrose biosynthesis and citric acid degradation genes for turning the fruit from sour to sweet.

Graphical AbstractFor image description, please refer to the figure legend and surrounding text.

## Linked entities

- **Chemicals:** ABA (PubChem CID 287291), sucrose (PubChem CID 5988), citric acid (PubChem CID 311)
- **Species:** Citrus reticulata (taxon 85571)

## Full-text entities

- **Genes:** bZIP5 (basic leucine-zipper 5) [NCBI Gene 824138] {aka AtbZIP5, basic leucine-zipper 5}, bZIP (basic leucine-zipper 8) [NCBI Gene 843221] {aka AtbZIP, T6L1.5, basic leucine-zipper 8}, HSFA6B (heat shock transcription factor A6B) [NCBI Gene 821854] {aka ARABIDOPSIS THALIANA HEAT SHOCK TRANSCRIPTION FACTOR A6B, AT-HSFA6B, heat shock transcription factor  A6B, heat shock transcription factor A6B}
- **Chemicals:** fructose (MESH:D005632), ACC (MESH:C023863), FAM (MESH:C031179), glucose (MESH:D005947), nitrogen (MESH:D009584), polyacrylamide (MESH:C016679), dithiothreitol (MESH:D004229), ABA (MESH:D000040), malate (MESH:C030298), sugar (MESH:D000073893), spermine (MESH:D013096), His (MESH:D006639), carbohydrate (MESH:D002241), Sucrose (MESH:D013395), poly (dI-dC) (MESH:C031156), IPTG (MESH:D007544), phenylmethylsulfonyl fluoride (MESH:D010664), Nonidet P-40 (MESH:C010615), tricarboxylic acid (MESH:D014233), Auxin (MESH:D007210), AOPI (-), Ethylene (MESH:C036216), DAP (MESH:C041756), Citric acid (MESH:D019343), carotenoids (MESH:D002338), cytokinins (MESH:D003583), EDTA (MESH:D004492), benzamidine (MESH:C032157), MgCl2 (MESH:D015636), DAPI (MESH:C007293)
- **Species:** Citrus sinensis (apfelsine, species) [taxon 2711], Citrus (genus) [taxon 2706], Salvia miltiorrhiza (Chinese salvia, species) [taxon 226208], Prunus persica (peach, species) [taxon 3760], Musa x paradisiaca (banana, species) [taxon 89151], Vitis vinifera (wine grape, species) [taxon 29760], Pyrus pyrifolia (Asian pear, species) [taxon 3767], Fragaria vesca (alpine strawberry, species) [taxon 57918], Solanum lycopersicum (tomato, species) [taxon 4081], Citrus x paradisi (grapefruit, species) [taxon 37656], Musa acuminata (banana, species) [taxon 4641], Malus domestica (apple, species) [taxon 3750], Saccharomyces cerevisiae (baker's yeast, species) [taxon 4932], Nicotiana benthamiana (species) [taxon 4100], Felis margarita (sand cat, species) [taxon 61378], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702], Mangifera indica (mango, species) [taxon 29780], Citrus reticulata (mandarin orange, species) [taxon 85571], Citrus unshiu (satsuma mandarin, species) [taxon 55188], Citrus japonica var. margarita (nagami kumquat, varietas) [taxon 85437], Pyrus x bretschneideri (bai li, species) [taxon 225117], Escherichia coli BL21 (strain) [taxon 511693], C. sinensis [taxon 128511], Fragaria x ananassa (strawberry, species) [taxon 3747], Agrobacterium tumefaciens (species) [taxon 358]

## Figures

9 figures with captions in the complete paper: https://tomesphere.com/paper/PMC13014036/full.md

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Source: https://tomesphere.com/paper/PMC13014036