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38 reviewsCarotenoids and flavonoids are two principal classes of secondary metabolites in tomato fruit, jointly contributing to pigmentation and nutritional quality. Here, we identified SlWRKY14 as a novel regulator of both thesetwo pathways. slwrky14 lines result in redder fruits enriched in carotenoids, whereas fruit-specific overexpression lines produce orange-yellow fruits with reduced carotenoids but elevated flavonoids. SlWRKY14directly binds to the promoter of carotenoid biosynthetic genes (SlDXS1 and SlPSY1) and represses theirtranscription by recruiting histone deacetylase SlHDA1, which functions in decreased histone acetylationto inhibit gene expression. Further genetic evidence verifies the role of the SlWRKY14-SlHDA1 module in suppressing carotenoid accumulation. In contrast, SlWRKY14 activates the flavonoid biosynthetic gene SlPALAto promote flavonoid accumulation. Our findings demonstrate SlWRKY14 as a regulatory node of carotenoidand flavonoid biosynthesis, providing a strategy for coloration regulation and quality improvement withoutpenalty of yield in tomato.