CsBBX20 Regulates Anthocyanin Biosynthesis in Citrus

doi:10.16420/j.issn.0513-353x.2026-0253

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (7): 2075-2091.doi: 10.16420/j.issn.0513-353x.2026-0253

• Genetic & Breeding · Germplasm Resources · Molecular Biology • Previous Articles Next Articles

CsBBX20 Regulates Anthocyanin Biosynthesis in Citrus

FU Jialing¹^,²^,^*(), WEI Yurong¹, SUN Juan², WEI Xing¹, DU Linzhu¹, HOU Jianqi¹, LIU Yuqi¹, XU Qiang²^,³^,^*()

¹ College of Agriculture， Guangxi University， Nanning 530004， China
² National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops;College of Horticulture and Forestry Sciences， Huazhong Agricultural University， Wuhan 430070， China
³ Hubei Hongshan Laboratory， Wuhan 430070， China

Received:2026-03-17 Revised:2026-05-20 Online:2026-07-25 Published:2026-07-23
Contact: FU Jialing, XU Qiang

Abstract

Abstract:

Anthocyanins are important secondary metabolites responsible for citrus fruit coloration，and their accumulation is regulated by environmental signals and multilayered regulatory networks. In this study，CsBBX20 was cloned from blood orange fruit. The open reading frame of CsBBX20 is 609 bp，encoding a protein of 202 amino acids. Sequence analysis showed that CsBBX20 contains two highly conserved B-box domains. Functional analysis revealed that anthocyanin accumulation was significantly increased in CsBBX20-overexpressing apple calli under light conditions，whereas only trace amounts of anthocyanins accumulated in wild-type calli. However，under dark conditions，neither CsBBX20-overexpressing lines nor wild-type calli accumulated anthocyanins. Further biochemical analyses demonstrated that CsBBX20 can directly bind to the promoters of CsF3’H，CsDFR，CsRuby1，and CsHY5，and significantly activate their expression. Moreover，CsBBX20 physically interacts with the light signaling regulator CsHY5，and their co-expression markedly enhances the activation of CsRuby1. Taken together，CsBBX20 directly activates key genes involved in anthocyanin biosynthesis and forms a complex with CsHY5 to coordinately regulate the expression of target genes，thereby promoting anthocyanin accumulation in citrus fruit under light conditions.

Key words: citrus, CsBBX20, anthocyanin, transcriptional regulation, CsHY5

FU Jialing, WEI Yurong, SUN Juan, WEI Xing, DU Linzhu, HOU Jianqi, LIU Yuqi, XU Qiang. CsBBX20 Regulates Anthocyanin Biosynthesis in Citrus[J]. Acta Horticulturae Sinica, 2026, 53(7): 2075-2091.

Figures/Tables 8

Table 1 Primers used in this study

用途 Purpose	引物名称 Primer name	引物序列（5′-3′） Primer sequence
qRT-PCR	qPCR-CsActin	F：CCGACCGTATGAGCAAGGAAA
		R：TTCCTGTGGACAATGGATGGA
	qPCR-CsBBX20	F：TCAGTCTTTTGCTCAGCCGATG
		R：AAGCAAAGCACGTCTCTCCTGA
	qPCR-MdActin	F：TGACCGAATGAGCAAGGAAATTACT
		R：TACTCAGCTTTGGCAATCCACATC
	qPCR-MdCHS	F：GGAGACAACTGGAGAAGGACTGGAA
		R：CGACATTGATACTGGTGTCTTC
	qPCR-MdF3’H	F：TGGAAGCTTGTGAGGACTGGGGT
		R：CTCCTCCGATGGCAAATCAAAGA
	qPCR-MdDFR	F：GATAGGGTTTGAGTTCAAGTA
		R：TCTCCTCAGCAGCCTCAGTTTTCT
	qPCR-MdANS	F：CCAAGTGAAGCGGGTTGTGCT
		R：CAAAGCAGGCGGACAGGAGTAGC
	qPCR-MdMYB1	F：TGCCTGGACTCGAGAGGAAGACA
		R：CCTGTTTCCCAAAAGCCTGTGAA
	qPCR-MdHY5	F：TGAAGAGGTTGTTGAGGAA
		R：TTCTGCAAAGTGGAAAGC
基因克隆 Gene cloning	CsBBX20-CDS	F：ATGAAGATTTGGTGTGATGTGTGTG
		R：TCACAAATATAAAACATTAGTA
	CsHY5-CDS	F：ATGCAAGAACAAGCAACGAGTTC
		R：TTAAAAGGACCCATCGGCAT
超表达载体 Overexpression vector	Adapter attB1	F：GGGGACAAGTTTGTACAAAAAAGCAGGCT
	Adapter attB2	R：GGGGACCACTTTGTACAAGAAAGCTGGGT
	CsBBX20-Adpter	F：AAAAAGCAGGCTCCATGAAGATTTGGTGTGATGTGTGTG
		R：AGAAAGCTGGGTTTCACAAATATAAAACATTAGTA
	CsHY5-Adpter	F：AAAAAGCAGGCTCCATGCAAGAACAAGCAACGAGTTC
		R：AGAAAGCTGGGTTTTAAAAGGACCCATCGGCAT
LUC实验 Dual-luciferase reporter assay	CsRuby1pro-LUC	F：GGTGGAGATCGAATTCCATGGTGTCAAAGTCGGTGGCCACAAGG
	CsRuby1pro-LUC	R：TGTTTTTGGCGTCTTCCATGGAACTCCTAAGGAATCCGCCAT
	CsHY5pro-LUC	F：GGTGGAGATCGAATTCCATGGCCACCAAAGATAATAGCAA
	CsHY5pro-LUC	R：TGTTTTTGGCGTCTTCCATGGTTCTTCTTCTCCTATTTCT
蛋白表达实验 Protein expression assay	CsBBX20-pET32a	F：GCTGATATCGGATCCGAATTCATGAAGATTTGGTGTGATGTGTGTG
蛋白表达实验 Protein expression assay	CsBBX20-pET32a	R：TGCGGCCGCAAGCTTGTCGACCAAATATAAAACATTAGTAGATG
EMSA实验 Electrophoretic mobility shift assay	CsRuby1-wt	F：GCATGAAGTGAGGAGCACGTATTATTATACAAGCAG
		R：CTGCTTGTATAATAATACGTGCTCCTCACTTCATGC
	CsF3’H-wt	F：GTGGTTACGTACTTACACGTACACCCCTACCTGCAT
		R：ATGCAGGTAGGGGTGTACGTGTAAGTACGTAACCAC
	CsDFR-wt	F：AAACGGTCAGTTGATCACGTGGAGGGTAATTGGAGC
		R：GCTCCAATTACCCTCCACGTGATCAACTGACCGTTT
	CsHY5-wt	F：AAACAAGAGAGAATACAAGTGAGCTTTATAAAGATG
		R：CATCTTTATAAAGCTCACTTGTATTCTCTCTTGTTT
Y2H实验 Yeast two-hybrid assay	CsBBX20-AD	F：GCCATGGAGGCCAGTGAATTCATGAAGATTTGGTGTGATGTGT
		R：ACGATTCATCTGCAGCTCGAGTCACAAATATAAAACATTAGTA
	CsHY5-BD	F：AGGACCTGCATATGGCCATGGATGCAAGAACAAGCAACGAGTTC
		R：TGCAGGTCGACGGATCCCCGGGTTAAAAGGACCCATCGGCATTTG
LCI实验 Luciferase complementation imaging assay	CsBBX20-nLUC	F：ACGGGGGACGAGCTCGGTACCATGAAGATTTGGTGTGATGT
		R：CGCGTACGAGATCTGGTCGACCAAATATAAAACATTAGTAG
	CsHY5-cLUC	F：TACGCGTCCCGGGGCGGTACCATGCAAGAACAAGCAACGAGTTC
		R：ACGAAAGCTCTGCAGGTCGACTTAAAAGGACCCATCGGCATTTG

Table 1 Primers used in this study

用途 Purpose	引物名称 Primer name	引物序列（5′-3′） Primer sequence
qRT-PCR	qPCR-CsActin	F：CCGACCGTATGAGCAAGGAAA
		R：TTCCTGTGGACAATGGATGGA
	qPCR-CsBBX20	F：TCAGTCTTTTGCTCAGCCGATG
		R：AAGCAAAGCACGTCTCTCCTGA
	qPCR-MdActin	F：TGACCGAATGAGCAAGGAAATTACT
		R：TACTCAGCTTTGGCAATCCACATC
	qPCR-MdCHS	F：GGAGACAACTGGAGAAGGACTGGAA
		R：CGACATTGATACTGGTGTCTTC
	qPCR-MdF3’H	F：TGGAAGCTTGTGAGGACTGGGGT
		R：CTCCTCCGATGGCAAATCAAAGA
	qPCR-MdDFR	F：GATAGGGTTTGAGTTCAAGTA
		R：TCTCCTCAGCAGCCTCAGTTTTCT
	qPCR-MdANS	F：CCAAGTGAAGCGGGTTGTGCT
		R：CAAAGCAGGCGGACAGGAGTAGC
	qPCR-MdMYB1	F：TGCCTGGACTCGAGAGGAAGACA
		R：CCTGTTTCCCAAAAGCCTGTGAA
	qPCR-MdHY5	F：TGAAGAGGTTGTTGAGGAA
		R：TTCTGCAAAGTGGAAAGC
基因克隆 Gene cloning	CsBBX20-CDS	F：ATGAAGATTTGGTGTGATGTGTGTG
		R：TCACAAATATAAAACATTAGTA
	CsHY5-CDS	F：ATGCAAGAACAAGCAACGAGTTC
		R：TTAAAAGGACCCATCGGCAT
超表达载体 Overexpression vector	Adapter attB1	F：GGGGACAAGTTTGTACAAAAAAGCAGGCT
	Adapter attB2	R：GGGGACCACTTTGTACAAGAAAGCTGGGT
	CsBBX20-Adpter	F：AAAAAGCAGGCTCCATGAAGATTTGGTGTGATGTGTGTG
		R：AGAAAGCTGGGTTTCACAAATATAAAACATTAGTA
	CsHY5-Adpter	F：AAAAAGCAGGCTCCATGCAAGAACAAGCAACGAGTTC
		R：AGAAAGCTGGGTTTTAAAAGGACCCATCGGCAT
LUC实验 Dual-luciferase reporter assay	CsRuby1pro-LUC	F：GGTGGAGATCGAATTCCATGGTGTCAAAGTCGGTGGCCACAAGG
	CsRuby1pro-LUC	R：TGTTTTTGGCGTCTTCCATGGAACTCCTAAGGAATCCGCCAT
	CsHY5pro-LUC	F：GGTGGAGATCGAATTCCATGGCCACCAAAGATAATAGCAA
	CsHY5pro-LUC	R：TGTTTTTGGCGTCTTCCATGGTTCTTCTTCTCCTATTTCT
蛋白表达实验 Protein expression assay	CsBBX20-pET32a	F：GCTGATATCGGATCCGAATTCATGAAGATTTGGTGTGATGTGTGTG
蛋白表达实验 Protein expression assay	CsBBX20-pET32a	R：TGCGGCCGCAAGCTTGTCGACCAAATATAAAACATTAGTAGATG
EMSA实验 Electrophoretic mobility shift assay	CsRuby1-wt	F：GCATGAAGTGAGGAGCACGTATTATTATACAAGCAG
		R：CTGCTTGTATAATAATACGTGCTCCTCACTTCATGC
	CsF3’H-wt	F：GTGGTTACGTACTTACACGTACACCCCTACCTGCAT
		R：ATGCAGGTAGGGGTGTACGTGTAAGTACGTAACCAC
	CsDFR-wt	F：AAACGGTCAGTTGATCACGTGGAGGGTAATTGGAGC
		R：GCTCCAATTACCCTCCACGTGATCAACTGACCGTTT
	CsHY5-wt	F：AAACAAGAGAGAATACAAGTGAGCTTTATAAAGATG
		R：CATCTTTATAAAGCTCACTTGTATTCTCTCTTGTTT
Y2H实验 Yeast two-hybrid assay	CsBBX20-AD	F：GCCATGGAGGCCAGTGAATTCATGAAGATTTGGTGTGATGTGT
		R：ACGATTCATCTGCAGCTCGAGTCACAAATATAAAACATTAGTA
	CsHY5-BD	F：AGGACCTGCATATGGCCATGGATGCAAGAACAAGCAACGAGTTC
		R：TGCAGGTCGACGGATCCCCGGGTTAAAAGGACCCATCGGCATTTG
LCI实验 Luciferase complementation imaging assay	CsBBX20-nLUC	F：ACGGGGGACGAGCTCGGTACCATGAAGATTTGGTGTGATGT
		R：CGCGTACGAGATCTGGTCGACCAAATATAAAACATTAGTAG
	CsHY5-cLUC	F：TACGCGTCCCGGGGCGGTACCATGCAAGAACAAGCAACGAGTTC
		R：ACGAAAGCTCTGCAGGTCGACTTAAAAGGACCCATCGGCATTTG

Fig. 1 Phylogenetic tree analysis of CsBBX20 At：Arabidopsis thaliana；Cs：Citrus sinensis；Fv：Fragaria vesca subsp. vesca；Cp：Carica papaya；Sl：Solanum lycopersicum；Pp：Prunus persica；Pa：Prunus avium；Pb：Pyrus × bretschneideri；Md：Malus × domestica. The same below

Fig. 2 Multiple sequence alignment of BBX20 protein sequences from different species Gray shading indicates amino acid sequence similarity greater than 75%

Fig. 3 Phenotypic characterization of CsBBX20-overexpressing apple calli A：Identification of CsBBX20-overexpressing apple calli，Bright-field images show the morphology of apple calli under natural light conditions，whereas GFP images represent calli observed under a fluorescence filter；B：Phenotypes of wild-type and CsBBX20-overexpressing apple calli under light and dark conditions；C，E：Expression levels of CsBBX20 in wild-type and CsBBX20-overexpressing apple calli under light（C）conditions and dark（E）conditions；D，F：Anthocyanin accumulation in wild-type and CsBBX20-overexpressing apple calli under both light（D）and dark（F）conditions. WT denotes the wild type，35S：CsBBX20 indicates CsBBX20-overexpressing lines，OE#1，OE#2，and OE#3 represent three independent CsBBX20-overexpressing lines. Student’s t-test，* P < 0.05，** P < 0.01，*** P < 0.001. The same below

Fig. 4 Expression of key genes in the anthocyanin biosynthetic pathway in CsBBX20-overexpressing apple calli under light（A）and dark（B）conditions

Fig. 5 Analysis of CsBBX20 regulation on the promoters of CsRuby1，CsF3’H and CsDFR A：Distribution of G-box cis-elements in the promoters of CsRuby1，CsF3’H，and CsDFR；B：Schematic representation of the constructs used in the dual-luciferase assay in Nicotiana benthamiana；C：Dual-luciferase assay validating the activation of CsRuby1，CsF3’H，and CsDFR promoters by CsBBX20，EV：pk7WG2D empty vector；D：EMSA showing the binding of CsBBX20-His to the G-box elements in the promoters of CsRuby1，CsF3’H，and CsDFR.“+”and“-”indicate the presence or absence of protein or probe，respectively. Wild-type probes were biotin-labeled（Bio-labeled）

Fig. 6 Analysis of the interaction between CsBBX20 and CsHY5 A：Yeast two-hybrid（Y2H）；B：Luciferase complementation imaging assay（LCI）（a indicated the nLUC + cLUC combination，b indicates the CsBBX20-nLUC + cLUC combination，c indicates the nLUC + CsHY5-cLUC combination，d indicates the CsBBX20-nLUC + CsHY5-cLUC combination）；C：Electrophoretic Mobility Shift Assay.“+”and“-”indicate the presence or absence of protein or probe，respectively，the WT probe was biotin-labeled，whereas the competitor（Competitor）and mutant probe（Mut probe）were unlabeled；D：Dual-Luciferase Reporter Assay in tobacco leaves；E：Tobacco LUC assay. 1 indicates the EV + CsRuby1pro combination，2 indicates EV + CsHY5 + CsRuby1pro，3 indicates EV + CsBBX20 + CsRuby1pro，4 indicates CsBBX20 + CsHY5 + CsRuby1pro

Fig. 7 Expression patterns of CsBBX20 under light and low-temperature treatments

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CsBBX20 Regulates Anthocyanin Biosynthesis in Citrus

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[4]	LI Weifen, ZHAO Dandan, LIU Fawan, FANG Chenggang, GUI Min, ZHANG Liqin, FENG Pengfei, CHEN Guofa, WU Zhen, LI Yinchuan, ZHOU Shan, DU Lei, ZHANG Ruihao, HU Huaran, ZHONG Qiuyue, WANG Renxiong, GAO Ye, DENG Zhengkui. A New Purple Leaf Mustard Cultivar‘Yunzijie 3’ [J]. Acta Horticulturae Sinica, 2026, 53(S1): 123-124.
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[9]	FU Qi, SHANG Junzhong, PEI Wenwen, LIU Guoliang, JING Weikun, ZHANG Hao, JIAN Hongying, QIU Xianqin, TANG Kaixue, YAN Huijun. Advances in the Biosynthetic Genes and Evolutionary Mechanisms of Floral Volatile Organic Compounds in Rosa hybrida [J]. Acta Horticulturae Sinica, 2026, 53(2): 481-495.
[10]	WEI Yonglu, YE Guangying, GAO Jie, JIN Jianpeng, LU Chuqiao, LI Jie, XIE Qie, GOU Yajun, ZHU Genfa, YANG Fengxi. Integrated Metabolome and Transcriptome Analysis of Floral Scent in Maxillaria tenuifolia [J]. Acta Horticulturae Sinica, 2026, 53(2): 538-556.
[11]	WANG Hong, WAN Yunqing, QI Xinai, SHENG Huizi, REN Yalin, LIU Mengyu, LIU Xiaofeng, ZHAO Xiaochun. Genome-Wide Identification of CYP71 Subfamily Genes in Citrus sinensis and Functional Analysis of CsCYP2 [J]. Acta Horticulturae Sinica, 2026, 53(1): 1-24.
[12]	KANG Jingbo, WEN Huan, WU Qun, AO Yijun, CHEN Jiajing, LIU Yuan, XU Juan. Comparative Analysis of Volatile Compounds in Seven Citrus junos Germplasms [J]. Acta Horticulturae Sinica, 2026, 53(1): 25-40.
[13]	FENG Huimin, LI Haibo, LEI Jianjun, ZHU Yunna. A New Mustard Cultivar‘Zimin Mustard’with High Anthocyanin [J]. Acta Horticulturae Sinica, 2025, 52(S2): 113-114.
[14]	LU Xin, REN Jie, DUAN Yaoyuan, XIE Kaidong, XIAO Gong’ao, XIA Zewen, WANG Rong, WU Xiaomeng, GUO Wenwu. Efficient in vivo Induction of Tetraploids from Three Mono-Embryonic Citrus Cultivars [J]. Acta Horticulturae Sinica, 2025, 52(9): 2291-2299.
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