STO represses anthocyanin synthesis and promotes flavonoid synthesis for photoprotection. Author Contributions: Shaoshan Li designed the project. Guizhen Lyu performed most of the experiments, and Dongbing Li and Hui Xiong participated in the experiments. Guizhen Lyu, Dongbing Li, and Chanjuan Ning analyzed the iTRAQ data. Langtao Xiao and Jian Hua Tong measured the plant hormones. Guizhen Lyu and Author Contributions: S.L. designed the project. G.L. performed most of the experiments, and D.L. and H.X. Dongbing Li wrote the draft of the manuscript, and Shaoshan Li revised the draft. All authors have read and participated in the experiments. G.L., D.L., and C.N. analyzed the iTRAQ data. L.X. and J.T. measured the plant approved the submitted vision of the manuscript. Funding acquisition and project administration were done by hormones. G.L. and D.L. wrote the draft of the manuscript, and S.L. revised the draft. Funding acquisition Shaoshan Li. and project administration were done by S.L. All authors have read and approved the submitted vision of the manuscript.Funding: This work was supportedby the National Natural Science Foundation of China (Grant No. 31670266 and 31570372), the Natural Science Foundation of Guangdong Province, China (Grant No. 2017A030313115 and Funding: This2w01o7rAk03w03a1s01s6u8p),pthoer ItnendovbaytiotnhePrNojeacttioofnGarlaNduaattuerSachloSoclioefnScoeutFhoCuhnindaaNtioornmoalfUCnhivienrasit(yG(2ra01n6tlkNxmo1.03),1670266 and 31570372),atnhdethNea LteuardainlgScSiceienncteistFsoPuronjedcattoifoGnuoafngGduonagngPdroovningceP arnodvitnheceG,uCanhgindoan(gGPreaanrltRNivoe.r2S0ch1o7lAar0F3u0n3d1e3d115 and 2017A03031016Sc8h)e, mthee( 2I0n12n)o. vation Project of Graduate School of South China Normal University (2016lkxm10), and the Leading Scientists Project of Guangdong Province and the Guangdong Pearl River Scholar Funded Scheme (2012).