Transcription factor CsHSFB2c suppresses CsTS1 and CsGS1 expression to reduce theanine biosynthesis under heat stress

doi:10.1016/j.jia.2025.12.027

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Transcription factor CsHSFB2c suppresses CsTS1 and CsGS1 expression to reduce theanine biosynthesis under heat stress

Qihong Zou¹, Bokun Zhou², Yilan Hu², Ping Li², Qi zhao², Hu Tang¹, Yujie Jiao¹, Xinzhuan Yao¹, Lin Chen², Litang Lu^{1, 2#}

¹College of Tea Science, Institute of Plant Health & Medicine, Guizhou University, Guiyang 550025, China

²The Key Laboratory of Plant Resources Conservation and Germplasm Innovation in Mountainous Region (Ministry of Education), College of Life Science, Guizhou University, Guiyang 550025, China

Hightlghts

l Heat stress led to the unique accumulation pattern of theanine and catechins in tea plants: Theanine content decreased significantly diminished with the increasing temperature (peaked at 20°C ), while catechins levels elevated with increasing temperature (peaked at 30°C).

l RNA-seq identified a class B heat shock transcription factor CsHSFB2c, whose expression was negatively correlated with theanine content.

l CsHSFB2c directly binds to the promoters of CsTS1 and CsGS1 and inhibits their transcription, thereby reducing theanine biosynthesis.

Abstract
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摘要

高温胁迫会降低茶树体内的茶氨酸含量，但其分子机制尚不明确。本研究通过设置温度梯度处理（20°C、25°C、30°C、35°C），旨在揭示高温胁迫对茶氨酸生物合成与积累的影响。研究发现，高温胁迫引发代谢重编程，其特征为茶氨酸含量降低而儿茶素含量升高。此外，高温胁迫上调了B类热休克转录因子基因CsHSFB2c的表达，同时显著抑制了茶氨酸生物合成关键基因CsTS1和CsGS1的转录。功能研究表明，沉默CsHSFB2c可提高茶氨酸含量，而过量表达该基因则显著降低茶氨酸水平。与上述结果一致，沉默CsHSFB2c会上调CsTS1和CsGS1的表达，而过表达CsHSFB2c则导致这两个基因的表达下调。酵母单杂交（Y1H）和双荧光素酶报告基因（Dual-LUC）实验表明，CsHSFB2c可直接结合CsTS1和CsGS1的启动子区域并抑制其表达。这些结果证明，CsHSFB2c通过直接抑制CsTS1和CsGS1的表达，介导了高温诱导的茶氨酸生物合成抑制。本研究为通过分子育种手段提高茶树抗热性及品质提供了理论依据。

Abstract

Theanine content in tea plants is reduced by heat stress, but its molecular mechanism is still unclear. In this study, a temperature gradient treatment (20°C, 25°C, 30°C, 35°C) was performed to unveil the effect of heat stress on biosynthesis and accumulation of theanine. It was found that heat stress triggered metabolic alterations characterized by reduced theanine and increased catechin levels. In addition, heat stress up-regulated the expression of class B heat shock transcription factor gene CsHSFB2c, while significantly suppressing the transcription of key theanine biosynthetic genes CsTS1 and CsGS1. Functional studies showed that silencing CsHSFB2c increased theanine content, while its overexpression significantly reduced theanine levels. Consistent with these changes, silencing CsHSFB2c up-regulated the expression of CsTS1 and CsGS1, while overexpression of CsHSFB2c resulted in their down-regulation. Yeast one-hybrid (Y1H) and dual-luciferase reporter gene (Dual-LUC) assays showed that CsHSFB2c directly binds to the promoters of CsTS1 and CsGS1 and inhibits their expression. These results demonstrate that CsHSFB2c mediates heat-induced suppression of theanine biosynthesis by directly inhibiting the expression of CsTS1 and CsGS1. This study provides a theoretical basis for improving heat resistance and quality of tea plants through molecular breeding.

Keywords: tea plant high temperature HSF glutamine synthetase theanine synthetase

Online: 18 December 2025

Fund:

This work was supported by Precise Breeding and Innovative Utilization of Guizhou Ancient Tea Germplasms (Qiankehe Major Project [2024]027), the Guizhou Province High-level Innovative Talents "Hundred" Level Talent Project (Qiankehe Platform Talent). GCC[2023]014; the Earmarked Fund for GZMARS-Tea (GZCYCYJSTX-03); the Research on the planting technology of China HUANENG photovoltaic tea garden (HNKJ2022-H135) and the introduced talent research fund project of Guizhou University (Gui Da Ren Ji He (2020) 55).

About author: #Correspondence Litang Lu, E-mail: ltlv@gzu.edu.cn

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Qihong Zou, Bokun Zhou, Yilan Hu, Ping Li, Qi zhao, Hu Tang, Yujie Jiao, Xinzhuan Yao, Lin Chen, Litang Lu. 2025. Transcription factor CsHSFB2c suppresses CsTS1 and CsGS1 expression to reduce theanine biosynthesis under heat stress. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2025.12.027

Bakery A, Vraggalas S, Shalha B, Chauchan H, Benhamed M, Fragkostefanakis S. 2024. Heat stress transcription factors as the central molecular rheostat to optimize plant survival and recovery from heat stress. New Phytologist, 244, 51-64.

Bharti K, von Koskull-Döring P, Bharti S, Kumar P, Tintschl-Körbitzer A, Treuter E, Nover L. 2004. Tomato heat stress transcription factor HsfB1 represents a novel type of general transcription coactivator with a histone-like motif interacting with the plant CREB binding protein ortholog HAC1. The Plant Cell, 16, 1521-1535.

Bian X H, Li W, Niu C F, Wei W, Hu Y, Han J Q, Lu X, Tao J J, Jin M, Qin H, Zhou B, Zhang W K, Ma B, Wang G D, Yu D Y, Lai Y C, Chen S Y, Zhang J S. 2020. A class B heat shock factor selected for during soybean domestication contributes to salt tolerance by promoting flavonoid biosynthesis. New Phytologist, 225, 268-283.

Chan E W C, Lim Y Y, Chew Y L. 2007. Antioxidant activity of Camellia sinensis leaves and tea from a lowland plantation in Malaysia. Food Chemistry, 102, 1214-1222.

Charng Y Y, Liu H C, Liu N Y, Chi W T, Wang C N, Chang S H, Wang T T. 2007. A heat-inducible transcription factor, HsfA2, is required for extension of acquired thermotolerance in Arabidopsis. Plant Physiology, 143, 251-262.

Chen F, He Y, Yao X Z, Zho B K, Tian S Y, Yin J, Lu L T. 2024. CsMOF1-guided regulation of drought-induced theanine biosynthesis in Camellia sinensis. International Journal of Biological Macromolecules, 268, 131725.

Chen H, Song Y J, Li H, Zaman S, Fan K, Ding Z T, Wang Y. 2023. Enhancing the adaptability of tea plants (Camellia sinensis L.) to high-temperature stress with small peptides and biosurfactants. Plants, 12, 2817.

Chen Z Z R, Yu Z X, Liu T T, Yao X Z, Zhang S Y, Hu Y L, Luo M Y, Wan Y, Lu L T. 2024. CsSPX3-CsPHL7-CsGS1/CsTS1 module mediated Pi-regulated negatively theanine biosynthesis in tea (Camellia sinensis). Horticulture Research, 11, uhae242.

Cheng S H, Fu X M, Wang X Q, Liao Y Y, Zeng L T, Dong F, Yang Z Y. 2017. Studies on the biochemical formation pathway of the amino acid L-theanine in tea (Camellia sinensis) and other plants. Journal of Agricultural and Food Chemistry, 65, 7210-7216.

Dai W D, Qi D D, Yang T, Lv H P, Guo L, Zhang Y, Zhu Y, Peng Q H, Xie D C, Tan J F. 2015. Nontargeted analysis using ultraperformance liquid chromatography–quadrupole time-of-flight mass spectrometry uncovers the effects of harvest season on the metabolites and taste quality of tea (Camellia sinensis L.). Journal of Agricultural and Food Chemistry, 63, 9869-9878.

Deng W W, Ogita S, Ashihara H. 2008. Biosynthesis of theanine (γ-ethylamino-l-glutamic acid) in seedlings of Camellia sinensis. Phytochemistry Letters, 1, 115-119.

Dong C X, Li F, Yang T Y, Feng L, Zhang S P, Li F D, Li W H, Xu G H, Bao S L, Wan X C Lucas W J, Zhang Z L. 2020. Theanine transporters identified in tea plants (Camellia sinensis L.). The Plant Journal, 101, 57-70.

Feng L, Gao M J, Hou R Y, Hu X Y, Zhang L, Wan X C, Wei S. 2014. Determination of quality constituents in the young leaves of albino tea cultivars. Food Chemistry, 155, 98-104.

Fragkostefanakis S, Röth S, Schleiff E, Schar K D. 2015. Prospects of engineering thermotolerance in crops through modulation of heat stress transcription factor and heat shock protein networks. Plant, Cell & Environment, 38, 1881-1895.

Fu X M, Chen Y Y, Mei X, Katsuno T, Kobayashi E, Dong F, Watanabe N, Yang Z Y. 2015. Regulation of formation of volatile compounds of tea (Camellia sinensis) leaves by single light wavelength. Scientific Reports, 5, 16858.

Fu X M, Liao Y Y, Cheng S H, Xu X, Grierson D, Yang Z Y. 2021. Nonaqueous fractionation and overexpression of fluorescent‐tagged enzymes reveals the subcellular sites of L-theanine biosynthesis in tea. Plant Biotechnology Journal, 19, 98-108.

Grentzmann G, Ingram J A, Kelly P J, Gestaland R F, Atkins J F. 1998. A dual-luciferase reporter system for studying recoding signals. Rna, 4, 479-486.

Hu Y L, Li P P, Yao X Z, He Y M, Tang H, Zhao Q, Lu L T. 2024. Zinc treatment of tea plants improves the synthesis of trihydroxylated catechins via regulation of the zinc-sensitive protein CsHIPP3. Journal of Agricultural and Food Chemistry, 72, 14887-14898.

Huang Y, An J, Sircar S, Bergis C, Lopes C D, He X N, Da C B, Tan F Q, Bazin J, Antunez-Sanchez J, Mammarella M F, Devani R S, Brik-Chaouche R, Bendahmane A, Frugier F, Xia C J, Rothan C, Probst A V, Mohamed Z, Bergounioux C, et al. 2023. HSFA1a modulates plant heat stress responses and alters the 3D chromatin organization of enhancer-promoter interactions. Nature Communications, 14, 469.

Ikeda M, Mitsuda N, Ohme-Takagi M. 2011. Arabidopsis HsfB1 and HsfB2b act as repressors of the expression of heat-inducible Hsfs but positively regulate the acquired thermotolerance. Plant Physiology, 157, 1243-1254.

Juneja L R, Chu D C, Okubo T, Nagato Y, Yokogoshi H. 1999. L-theanine — a unique amino acid of green tea and its relaxation effect in humans. Trends in Food Science and Technology, 10, 199-204.

Li B J, Jiang S M, Gao L, Wang W H, Luo H Z, Dong Y N, Gao Z H, Zheng S Z, Liu X Y, Tang W Q. 2024. Heat shock factor A1s are required for phytochrome-interacting factor 4-mediated thermomorphogenesis in Arabidopsis. Journal of Integrative Plant Biology, 66, 20-35.

Li F, Dong C X, Yang T Y, Ma J Z, Zhang S P, Wei C L, Wan X C , Zhang Z L. 2019. Seasonal theanine accumulation and related gene expression in the roots and leaf buds of tea plants (Camellia sinensis L.). Frontiers in Plant Science, 10, 1397.

Li G D, Li Y, Yao X Z, Lu L T. 2022. Establishment of a virus-induced gene-silencing (VIGS) system in tea plant and its use in the functional analysis of CsTCS1. International Journal of Molecular Sciences, 24, 392.

Li H, Guo L H, Yan M L, Hu J, Lin Q Q, Wang P, Wang M L, Zhao H, Wang Y, Ni D J. 2022. A rapid and efficient transient expression system for gene function and subcellular localization studies in the tea plant (Camellia sinensis) leaves. Scientia Horticulturae, 297, 110927.

Li X, Ahammed G J, Li Z X, Zhang L, Wei J P, Shen C, Yan P, Zhang L P, Han W Y. 2016. Brassinosteroids improve quality of summer tea (Camellia sinensis L.) by balancing biosynthesis of polyphenols and amino acids. Frontiers in Plant Science, 7, 1304.

Li X, Wei J P, Ahammed G J, Zhang L, Li Y, Yan P, Zhang L P, Han W Y. 2018. Brassinosteroids attenuate moderate high temperature-caused decline in tea quality by enhancing theanine biosynthesis in Camellia sinensis L. Frontiers in Plant Science, 9, 1016.

Lin N, Liu X Y, Zhu W F, Cheng X, Wang X H, Wan X C, Liu L L. 2021. Ambient ultraviolet B signal modulates tea flavor characteristics via shifting a metabolic flux in flavonoid biosynthesis. Journal of Agricultural and Food Chemistry, 69, 3401-3414.

Liu J W, Zhang Q F, Liu M Y, Ma L F, Shi Y Z, Ruan J Y. 2016. Metabolomic analyses reveal distinct change of metabolites and quality of green tea during the short duration of a single spring season. Journal of Agricultural and Food Chemistry, 64, 3302-3309.

Liu Z W, Li H, Wang W L, Wu Z J, Cui X, Zhuang J. 2017. CsGOGAT is important in dynamic changes of theanine content in postharvest tea plant leaves under different temperature and shading spreadings. Journal of Agricultural and Food Chemistry, 65, 9693-9702.

Lu L T, Chen H F, Wang X J, Zhao Y C, Yao X Z, Xiong B, Deng Y L, Zhao D J. 2021. Genome-level diversification of eight ancient tea populations in the Guizhou and Yunnan regions identifies candidate genes for core agronomic traits. Horticulture Research, 8,190-190.

Lü X P, Shao K Z, Xu J Y, Li J L, Ren W, Chen J, Zhao L Y, Zhao Q, Zhang J L. 2022. A heat shock transcription factor gene (HaHSFA1) from a desert shrub, Haloxylon ammodendron, elevates salt tolerance in Arabidopsis thaliana. Environmental and Experimental Botany, 201, 104954.

Raturi V, Zinta G. 2024. HSFA1 heat shock factors integrate warm temperature and heat signals in plants. Trends in Plant Science, 29, 1165-1167.

Ren T Y, Zheng P C, Zhang K X, Liao J R, Xiong F, Shen Q, Ma Y C, Fang W P, Zhu X J. 2021. Effects of GABA on the polyphenol accumulation and antioxidant activities in tea plants (Camellia sinensis L.) under heat-stress conditions. Plant Physiology and Biochemistry, 159, 363-371.

Scharf K D, Berberich T, Ebersberger I, Nover L. 2012. The plant heat stress transcription factor (Hsf) family: structure, function and evolution. Biochimica et Biophysica Acta (BBA) - Gene Regulatory Mechanisms, 1819, 104-119.

Tan W R, Chen J H, Yue X L, Chai S L, Liu W, Li C L, Yang F, Gao Y F, Gutiérrez Rodríguez L, Resco de Dios V, Zhang D W, Yao Y N. 2023. The heat response regulators HSFA1s promote Arabidopsis thermomorphogenesis via stabilizing PIF4 during the day. Science Advances, 9, eadh1738.

Tsushida T, Takeo T. 1984. Ethylamine content of fresh tea shoots and made tea determined by high performance liquid chromatography. Journal of the Science of Food and Agriculture, 35, 77-83.

Wang H R, Feng M, Jiang Y J, Du D J, Dong C Q, Zhang Z H, Wang W X, Liu J, Liu X Q, Li S F, Chen Y M, Guo W L, Xin M M, Yao Y Y, Ni Z F, Sun Q X, Peng H R, Liu J. 2023. Thermosensitive SUMOylation of TaHsfA1 defines a dynamic ON/OFF molecular switch for the heat stress response in wheat. The Plant Cell, 35, 3889-3910.

Wei C L, Yang H, Wang S B, Zhao J, Liu C, Gao L P, Xia E H, Lu Y, Tai Y L, She G B, Sun J, Cao H S, Tong W, Gao Q, Li Y Y, Deng W W, Jiang X L, Wang W Z, Chen Q, Zhang S H, et al. 2018. Draft genome sequence of Camellia sinensis var. sinensis provides insights into the evolution of the tea genome and tea quality. Proceedings of the National Academy of Sciences, 115, E4151-E4158.

Wen B B, Luo Y, Liu D M, Zhang X N, Peng Z, Wang K B, Li J, Huang J A, Liu Z H. 2020. The R2R3-MYB transcription factor CsMYB73 negatively regulates l-Theanine biosynthesis in tea plants (Camellia sinensis L.). Plant Science, 298, 110546.

Wu W, Wang M M, Gong H, Liu X F, Guo D L, Sun N J, Huang J W, Zhu Q G, Chen K S, Yin X R. 2020. High CO₂/hypoxia-induced softening of persimmon fruit is modulated by DkERF8/16 and DkNAC9 complexes. Journal of Experimental Botany, 71, 2690-2700.

Xu W P, Song Q S, Li D X, Wan X C. 2012. Discrimination of the production season of Chinese green tea by chemical analysis in combination with supervised pattern recognition. Journal of Agricultural and Food Chemistry, 60, 7064-7070.

Yuan Z K D, Tian X Y, Yao X Z, Ye S Y, Li P, Tang H, Jiao Y J, Zhao Q, Lu L T. 2025. The CsGT1A-CsSCPL11-IA module positively regulates galloylated catechin biosynthesis in tea plants. Industrial Crops and Products, 227, 120859.

Zhang S P, Lin S J, Jin R P, Zhang Y N, Zhang X X, Zhu B Y, Luo Y Y, Ham B K, Chen J Y, Yang T Y, Wan X C, Zhang Z L. 2025. CATIONIC AMINO ACID TRANSPORTER 1 modulates amino acid distribution between stem and leaf in new shoots: A case study of theanine distribution in tea plants (Camellia sinensis). Plant Physiology, 198, kiaf255.

Zhang X Y, Xu W L, Ni D J, Wang M L, Guo G Y. 2020. Genome-wide characterization of tea plant (Camellia sinensis) Hsf transcription factor family and role of CsHsfA2 in heat tolerance. BMC Plant Biology, 20, 1-17.

Zhang Y R, Li P H, She G H, Xu Y J, Peng A Q, Wan X C, Zhao J. 2021. Molecular basis of the distinct metabolic features in shoot tips and roots of tea plants (Camellia sinensis): characterization of MYB regulator for root theanine synthesis. Journal of Agricultural and Food Chemistry, 69, 3415-3429.

Zhu Q F, Liu L J, Lu X F, Du X X, Xiang P, Cheng B S, Tan M, Huang J X, Wu L J, Kong W L, Shi Y T, Wu L Y, Lin J K. 2023. The biosynthesis of EGCG, theanine and caffeine in response to temperature is mediated by hormone signal transduction factors in tea plant (Camellia sinensis L.). Frontiers in Plant Science, 14, 1149182.

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