MdWRKY33 and MdWRKY127 Regulate Sugar and Organic Acid Metabolism to Improve the Sugar-Acid Ratio in Apple Fruit

doi:10.16420/j.issn.0513-353x.2026-0120

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (7): 1991-2003.doi: 10.16420/j.issn.0513-353x.2026-0120

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

MdWRKY33 and MdWRKY127 Regulate Sugar and Organic Acid Metabolism to Improve the Sugar-Acid Ratio in Apple Fruit

SU Anqi¹, ZHUANG Yingyu¹, DU Lianda¹, PEI Feifei², CHEN Ze¹, ZHAO Tingting¹^,^**(), HU Dagang¹^,^**()

¹ College of Horticultural Science and Engineering， Shandong Agricultural University，Tai?an， Shandong 271018， China
² Shandong Institute of Commerce and Technology， Ji?nan 250103， China

Received:2026-01-28 Revised:2026-06-12 Online:2026-07-25 Published:2026-07-23
Contact: ZHAO Tingting, HU Dagang

Abstract

Abstract:

Transcriptome analysis of‘Greensleeves’apple（Malus × domestica Borkh.）fruit during developmental stages identified two WRKY transcription factors，MdWRKY33 and MdWRKY127，which were specifically highly expressed at the ripening stage. To clarify the biological functions of these two WRKY transcription factors，they were cloned and subjected to bioinformatics analysis. Phylogenetic

analysis revealed that the MdWRKY33 protein shares close phylogenetic relationships with its homologous proteins from wild apple relative Siberian crabapple（Malus baccatavar. baccata）and pear（Pyrus× bretschneideri），while the MdWRKY127 protein is closely related to an incompletely annotated protein in apple. Subcellular localization analysis indicated that both MdWRKY33 and MdWRKY127 proteins are located in the nucleus，exhibiting the typical subcellular localization characteristics of WRKY transcription factors. To verify their functions，overexpression experiments were carried out using apple calli as the material. The results showed that overexpressing either MdWRKY33or MdWRKY127gene significantly reduced the content of malic acid and citric acid in the calli，while increasing the accumulation levels of fructose and sucrose，resulting in a higher sugar-acid ratio in the calli. Further transient injection experiments in apple fruits showed that the overexpression results were consistent with those in the calli，whereas silencing either gene led to a decrease in the sugar-acid ratio in the fruit. These results indicate that both MdWRKY33 and MdWRKY127 are involved in regulating sugar and acid metabolism in apple fruit，thereby effectively improving the flavor of apple fruit.

Key words: apple, WRKY, fruit flavor, soluble sugars, organic acids, sugar-acid ratio, overexpression, calli

SU Anqi, ZHUANG Yingyu, DU Lianda, PEI Feifei, CHEN Ze, ZHAO Tingting, HU Dagang. MdWRKY33 and MdWRKY127 Regulate Sugar and Organic Acid Metabolism to Improve the Sugar-Acid Ratio in Apple Fruit[J]. Acta Horticulturae Sinica, 2026, 53(7): 1991-2003.

Figures/Tables 10

Table 1 Primers used in this study

引物用途Primer application	引物名称 Primer name	引物序列（5′-3′）Primer sequence
基因克隆 Gene cloning	MdWRKY33-F	ATGACGGTGGAGCTGATGGAAT
	MdWRKY33-R	CGTTGACTCGTACACCAACCCC
	MdWRKY127-F	ATGGCTATAGATCTAGTTGGCTTCT
	MdWRKY127-R	GGACGACTGGAAAACAAGGCC
亚细胞定位及愈伤过表达 Subcellular localization and callus overexpression	MdWRKY33-3HA-F	GGTACCCGGGGATCCATGACGGTGGAGCTGATGGAAT
	MdWRKY33-3HA-R	CACCATGGTGTCGACCGTTGACTCGTACACCAACCCC
	MdWRKY127-3HA-F	GGTACCCGGGGATCCATGGCTATAGATCTAGTTGGCTTCT
	MdWRKY127-3HA-R	CACCATGGTGTCGACGGACGACTGGAAAACAAGGCC
实时荧光定量PCR qRT-PCR	qMdWRKY33-F	GAGATGGCAGCGTTTCGAAC
	qMdWRKY33-R	CCGTTGAGAGAGGAGGCTTC
	qMdWRKY127-F	ATCGATTTCTGTGCCTACGC
	qMdWRKY127-R	ACTCTGATCGTCCTCTTAACCT
内参基因 Reference gene	18S-F	ACACGGGGAGGTAGTGACAA
内参基因 Reference gene	18S-R	CCTCCAATGGATCCTCGTTA
基因过表达 Gene overexpression	MdWRKY33-pIR-F	TCTGAATTCGTCGACAAGCTTATGACGGTGGAGCTGATGGAAT
	MdWRKY33-pIR-R	CCCCCACACGTGTGGTCTAGACGTTGACTCGTACACCAACCCC
	MdWRKY127-pIR-F	TCTGAATTCGTCGACAAGCTTATGGCTATAGATCTAGTTGGCTTCT
	MdWRKY127-pIR-R	CCCCCACACGTGTGGTCTAGAGGACGACTGGAAAACAAGGCC
基因沉默 Gene silencing	MdWRKY33-TRV-F	AGAAGGCCTCCATGGGGATCCATGACGGTGGAGCTGATGGAATT
	MdWRKY33-TRV-R	TGTCTTCGGGACATGCCCGGGGGAGACGGTGTGATCGGTGATG
	MdWRKY127-TRV-F	AGAAGGCCTCCATGGGGATCCATGGCTATAGATCTAGTTGGCTTCT
	MdWRKY127-TRV-R	TGTCTTCGGGACATGCCCGGGGTGCTTGAACTTGGTGACGGTGA

Fig. 1 Heat map of MdWRKY33 and MdWRKY127 gene expression in‘Greensleeves’apple fruits at 16，41，70，94，and 128 days after flowering n = 3

Fig. 2 Gene structural analysis of MdWRKY33 and MdWRKY127 in apple The black part represents the exon，the black line（intron）represents the intron，and the white part represents the untranslated region

Fig. 3 Phylogenetic tree of MdWRKY33 and MdWRKY127 proteins in apple

Fig. 4 Multiple sequence alignment of apple MdWRKY33，MdWRKY127 proteins and their homologous proteins from other species The red box shows the WRKY domain

Fig. 5 Subcellular localization of apple MdWRKY33 and MdWRKY127 proteins

Fig. 6 Phenotypes（A），gene expression levels（B）and molecular identification（C）of apple MdWRKY33 and MdWRKY127 transgenic calli WT represents wild-type control. Duncan’s multiple range test was used for significance analysis，and different lowercase letters indicate significant differences at P < 0.05.The same below

Fig. 7 Sugar and organic acid contents in apple transgenic calli overexpressing MdWRKY33（A）and MdWRKY127（B）

Fig. 8 Phenotype and identification of transiently injected MdWRKY33（A）and MdWRKY127（B）in apples 1：pIR + TRV（Control 1）；2：MdWRKY33-pIR + TRV；3：pIR + MdWRKY33-TRV；4：pIR + TRV（Control 2）；5：MdWRKY127-pIR + TRV；6：pIR + MdWRKY127-TRV. The same below

Fig. 9 Sugar and acid content of transiently injected MdWRKY33（A）and MdWRKY127（B）in apples

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MdWRKY33 and MdWRKY127 Regulate Sugar and Organic Acid Metabolism to Improve the Sugar-Acid Ratio in Apple Fruit

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