Tea-Derived Polyphenols Enhance Drought Resistance of Tea Plants (Camellia sinensis) by Alleviating Jasmonate-Isoleucine Pathway and Flavonoid Metabolism Flow.

Extreme drought weather has occurred frequently in recent years, resulting in serious yield loss in tea plantations. The study of drought in tea plantations is becoming more and more intensive, but there are fewer studies on drought-resistant measures applied in actual production. Therefore, in this study, we investigated the effect of exogenous tea polyphenols on the drought resistance of tea plant by pouring 100 mg·L-1 of exogenous tea polyphenols into the root under drought. The exogenous tea polyphenols were able to promote the closure of stomata and reduce water loss from leaves under drought stress. Drought-induced malondialdehyde (MDA) accumulation in tea leaves and roots was also significantly reduced by exogenous tea polyphenols. Combined transcriptomic and metabolomic analyses showed that exogenous tea polyphenols regulated the abnormal responses of photosynthetic and energy metabolism in leaves under drought conditions and alleviated sphingolipid metabolism, arginine metabolism, and glutathione metabolism in the root system, which enhanced the drought resistance of tea seedlings. Exogenous tea polyphenols induced jasmonic acid-isoleucine (JA-ILE) accumulation in the root system, and the jasmonic acid-isoleucine synthetase gene (TEA028623), jasmonic acid ZIM structural domain proteins (JAMs) synthesis genes (novel.22237, TEA001821), and the transcription factor MYC2 (TEA014288, TEA005840) were significantly up-regulated. Meanwhile, the flavonoid metabolic flow was significantly altered in the root; for example, the content of EGCG, ECG, and EGC was significantly increased. Thus, exogenous tea polyphenols enhance the drought resistance of tea plants through multiple pathways.

Flavonoid metabolites in tea plant (Camellia sinensis) stress response: Insights from bibliometric analysis.

In the context of global climate change, tea plants are at risk from elevating environmental stress factors. Coping with this problem relies upon the understanding of tea plant stress response and its underlying mechanisms. Over the past two decades, research in this field has prospered with the contributions of scientists worldwide. Aiming in providing a comprehensive perspective of the research field related to tea plant stress response, we present a bibliometric analysis of the this area. Our results demonstrate the most studied stresses, global contribution, authorship and collaboration, and trending research topics. We highlight the importance of flavonoid metabolites in tea plant stress response, particularly their role in maintaining redox homeostasis, yield, and adjusting tea quality under stress conditions. Further research on the flavonoid response under various stress conditions can promote the development of cultivation measures, thereby improving stress resistance and tea quality.

Topics and trends in fresh tea (Camellia sinensis) leaf research: A comprehensive bibliometric study.

Tea plant (Camellia sinensis) is a widely cultivated cash crop and tea is a favorite functional food in the world. Fresh tea leaves (FTLs) play a critical role in bridging the two fields closely related to tea cultivation and tea processing, those are, tea plant biology and tea biochemistry. To provide a comprehensive overview of the development stages, authorship collaboration, research topics, and hotspots and their temporal evolution trends in the field of FTLs research, we conducted a bibliometric analysis, based on 971 publications on FTLs-related research published during 2001-2021 from Web of Science Core Collection. CiteSpace, R package Bibliometrix, and VOSviewer were employed in this research. The results revealed that the development history can be roughly divided into three stages, namely initial stage, slow development stage and rapid development stage. Journal of Agricultural & Food Chemistry published most articles in this field, while Frontiers in Plant Science held the highest total citations and h-index. The most influential country, institution, and author in this field was identified as China, the Chinese Academy of Agricultural Sciences, and Xiaochun Wan, respectively. FTLs-related research can be categorized into three main topics: the regulation mechanism of key genes, the metabolism and features of essential compounds, and tea plants' growth and stress responses. The most concerning hotspots are the application of advanced technologies, essential metabolites, leaf color variants, and effective cultivation treatments. There has been a shift from basic biochemical and enzymatic studies to studies of molecular mechanisms that depend on multi-omics technologies. We also discussed the future development in this field. This study provides a comprehensive summary of the research field, making it easier for researchers to be informed about its development history, status, and trends.

Effects of Preharvest Shading on Dynamic Changes in Metabolites, Gene Expression, and Enzyme Activity of Three Tea Types during Processing.

Preharvest shading significantly influences tea flavor. However, little attention has been given to the mechanism of shading on metabolites, genes, and enzymes in the processing of different tea types. Our study identified 1028 nonvolatile metabolites covering 10 subclasses using a widely targeted metabolome. The results show that shading had a greater effect on the compositions of amino acids, flavonoids, and theaflavins in tea leaves. The combined transcriptomics and enzyme activity analysis results indicate that the upregulated expression of asparagine, aspartate, and tryptophan synthesis genes and proteolytic enzymes promoted the accumulation of amino acids. The downregulated enzyme genes resulted in the reduction of nongalloylated catechins and flavonoid glycosides. Simultaneously, the accumulation of TFs in shaded tea was due to the enhanced enzymatic activities of polyphenol oxidase and peroxidase during processing. Theaflavin-3-3'-di-O-gallate was also significantly positively correlated with the antioxidant and hypoglycemic activities of shaded tea. The results contribute to a better understanding of how preharvest treatments influence summer tea quality.

Carbon and Nitrogen Metabolism Are Jointly Regulated During Shading in Roots and Leaves of Camellia Sinensis.

Numerous studies have shown that plant shading can promote the quality of green tea. However, the association of shading with metabolic regulation in tea leaves and roots remains unelucidated. Here, the metabolic profiling of two tea cultivars ("Xiangfeicui" and "Jinxuan") in response to shading and relighting periods during the summer season was performed using non-targeted metabolomics methods. The metabolic pathway analyses revealed that long-term shading remarkably inhibit the sugar metabolism such as glycolysis, galactose metabolism, and pentose phosphate pathway in the leaves and roots of "Xiangfeicui," and "Jinxuan" were more sensitive to light recovery changes. The lipid metabolism in the leaves and roots of "Xiangfeicui" was promoted by short-term shading, while it was inhibited by long-term shading. In addition, the intensity of the flavonoid metabolites in the leaves and roots of "Jinxuan" were upregulated with a trend of rising first and then decreasing under shading, and five flavonoid synthesis genes showed the same trend (F3H, F3'5'H, DFR, ANS, and ANR). Simultaneously, the amino acids of the nitrogen metabolism in the leaves and roots of the two cultivars were significantly promoted by long-term shading, while the purine and caffeine metabolism was inhibited in the leaves of "Xiangfeicui." Interestingly, CsGS1.1 and CsTSI, amino acid synthase genes was upregulated in the leaves and roots of two cultivars. These results indicated that shading could participate in carbon and nitrogen metabolic regulation of both leaf and root, and root metabolism could have a positive association with leaf metabolism to promote the shaded tea quality.

Transcriptomic analyses reveal variegation-induced metabolic changes leading to high L-theanine levels in albino sectors of variegated tea (Camellia sinensis).

Camellia sinensis cv. 'Yanling Huayecha' (YHC) is an albino-green chimaeric tea mutant with stable genetic traits. Here, we analysed the cell ultrastructure, photosynthetic pigments, amino acids, and transcriptomes of the albino, mosaic, and green zones of YHC. Well-organized thylakoids were found in chloroplasts in mesophyll cells of the green zone but not the albino zone. The albino zone of the leaves contained almost no photosynthetic pigment. However, the levels of total amino acids and theanine were higher in the albino zone than in the mosaic and green zones. A transcriptomic analysis showed that carbon metabolism, nitrogen metabolism and amino acid biosynthesis showed differences among the different zones. Metabolite and transcriptomic analyses revealed that (1) downregulation of CsPPOX1 and damage to thylakoids in the albino zone may block chlorophyll synthesis; (2) downregulation of CsLHCB6, CsFdC2 and CsSCY1 influences chloroplast biogenesis and thylakoid membrane formation, which may contribute to the appearance of variegated tea leaves; and (3) tea plant variegation disrupts the balance between carbon and nitrogen metabolism and promotes the accumulation of amino acids, and upregulation of CsTSⅠ and CsAlaDC may enhance L-theanine synthesis. In summary, our study provides a theoretical basis and valuable insights for elucidating the molecular mechanisms and promoting the economic utilization of variegation in tea.

Disruption of Photomorphogenesis Leads to Abnormal Chloroplast Development and Leaf Variegation in Camellia sinensis.

Camellia sinensis cv. 'Yanlingyinbiancha' is a leaf-variegated mutant with stable genetic traits. The current study aimed to reveal the differences between its albino and green tissues, and the molecular mechanism underlying the variegation. Anatomic analysis showed the chloroplasts of albino tissues to have no intact lamellar structure. Photosynthetic pigment in albino tissues was significantly lower than that in green tissues, whereas all catechin components were more abundant in the former. Transcriptome analysis revealed most differentially expressed genes involved in the biosynthesis of photosynthetic pigment, photosynthesis, and energy metabolism to be downregulated in albino tissues while most of those participating in flavonoid metabolism were upregulated. In addition, it was found cryptochrome 1 (CRY1) and phytochrome B (PHYB) genes that encode blue and red light photoreceptors to be downregulated. These photoreceptors mediate chloroplast protein gene expression, chloroplast protein import and photosynthetic pigment biosynthesis. Simultaneously, SUS gene, which was upregulated in albino tissues, encodes sucrose synthase considered a biochemical marker for sink strength. Collectively, we arrived to the following conclusions: (1) repression of the biosynthesis of photosynthetic pigment causes albinism; (2) destruction of photoreceptors in albino tissues suppresses photomorphogenesis, leading to abnormal chloroplast development; (3) albino tissues receive sucrose from the green tissues and decompose their own storage substances to obtain the energy needed for survival; and (4) UV-B signal and brassinosteroids promote flavonoid biosynthesis.

Research progress on the response of tea catechins to drought stress.

Drought stress (DS) is the most important abiotic stress affecting yield and quality of tea worldwide. DS causes oxidative stress to cells due to the accumulation of reactive oxygen species (ROS). As non-enzymatic antioxidants, tea catechins can scavenge excess ROS in response to DS. Further, catechin accumulation contributes to the formation of oxidative polymerization products (e.g. theaflavins and thearubigins) that improve the quality of black tea. However, there are no systematic reports on the response of tea catechins to DS. First, we reviewed the available literature on the response of tea plants to DS. Second, we summarized the current knowledge of ROS production in tea leaves under DS and typical antioxidant response mechanisms. Third, we conducted a detailed review of the changes in catechin levels in tea under different drought conditions. We found that the total amounts of catechin and o-quinone increased under DS conditions. We propose that the possible mechanisms underlying tea catechin accumulation under DS conditions include (i) autotrophic formation of o-quinone, (ii) polymerization of proanthocyanidins that directly scavenge excess ROS, and (iii) formation of metal ion complexes and by influencing the antioxidant systems that indirectly eliminate excess ROS. Finally, we discuss ways of potentially improving black tea quality using drought before picking in the summer/fall dry season. In summary, we mainly discuss the antioxidant mechanisms of tea catechins under DS and the possibility of using drought to improve black tea quality. Our review provides a theoretical basis for the production of high-quality black tea under DS conditions. © 2021 Society of Chemical Industry.

Survival strategies based on the hydraulic vulnerability segmentation hypothesis, for the tea plant [Camellia sinensis(L.) O. Kuntze] in long-term drought stress condition.

Tea plants are important economic perennial crops that can be negatively impacted by drought stress (DS). However, their survival strategies in long-term DS conditions and the accumulation and influence of metabolites and mineral elements (MEs) in their organs, when facing hydraulic vulnerability segmentation, require further investigation. The MEs and metabolites in the leaf, stem, and root after long-term DS (20 d) were examined here, using inductively coupled plasma optical emission spectrometry (ICP-OES) and liquid chromatograph-mass spectrometry (LC-MS). The accumulation patterns of 116 differentially accumulated metabolites (DAMs) and nine MEs were considerably affected in all organs. The concentration of all MEs varied significantly in at least one organ, while the K and Ca levels were markedly altered in all three. Most DAM levels increased in the stem but decreased in the root and leaf, implying that vulnerability segmentation may occur with long-term DS. The typical nitrogen- and carbon-compound levels similarly increased in the stem and decreased in the leaf and root, as the plant might respond to long-term DS by stabilizing respiration, promoting nitrogen recycling, and free radical scavenging. Correlation analysis showed several possible DAM-ME interactions and an association between Mn and flavonoids. Thus, survival strategies under long-term DS included sacrificing distal/vulnerable organs and accumulating function-specialized metabolites and MEs to mitigate drought-induced oxidative damage. This is the first study that reports substance fluctuations after long-term DS in different organs of plants, and highlights the need to use whole plants to fully comprehend stress response strategies.

Physiological and biochemical responses of tea seedlings (Camellia sinensis) to simulated acid rain conditions.

Tea (Camellia sinensis), widely planted in the south of China, and often exposed to acid rain. However, research concerning the impacts of acid rain on physiology and biochemistry of tea plants is still scarce. In this study, we investigated the influence of simulated acid rain (SAR) on plant height, root length, photosynthetic pigment, Fv/Fm, proline, malondialdehyde, antioxidant enzyme activity, total nitrogen, caffeine, catechins, and free amino acids. Our results showed that SAR at pH 4.5 did not hinder plant development because growth characteristics, photosynthesis, and ascorbate peroxidase and catalase activities did not decrease at this pH compared to those at the other investigated pH values. However, at pH 3.5 and pH 2.5, the activities of antioxidase and concentrations of malondialdehyde and proline increased significantly in response to the decrease of photosynthetic pigments and Fv/Fm. In addition, the increase in acidity increased total nitrogen, certain amino acid content (theanine, cysteine), and decreased catechin and caffeine contents, resulting in an imbalance of the carbon and nitrogen metabolisms. Our results indicated that SAR at pH 3.5 and pH 2.5 could restrict photosynthesis and the antioxidant defense system, causing metabolic disorders and ultimately affecting plant development and growth, but SAR at pH 4.5 had no toxic effects on tea seedlings when no other stress factors are involved.

Exogenous indole acetic acid alleviates Cd toxicity in tea (Camellia sinensis).

Cadmium (Cd), a toxic heavy metal, restrains the growth and development of plants and threatens global food safety. Many studies on the alleviation of heavy metal toxicity by exogenous phytohormones have emerged, but reports on tea (Camellia sinensis) are still scarce. In this study, the effects of indole acetic acid (IAA) (2 µM and 10 µM) on Cd uptake and on the physiological and biochemical characteristics of the 'Xiangfeicui' tea cultivar were investigated for the first time. The order of Cd accumulation in tea seedlings was root > stem > mature leaf > tender leaf. Under Cd stress (30 mg kg-1), photosynthetic pigment levels, antioxidant enzyme activity, root vigor, root IAA content, and the levels of most metabolites (including caffeine, soluble sugar, total amino acids, some amino acid components, and most catechins) were significantly reduced, while levels of malondialdehyde, proline, epicatechin, and some amino acids increased. We therefore propose that by reducing Cd accumulation, exogenous IAA can lessen the adverse effects of Cd on the physiology and biochemistry of tea seedlings, promoting the growth of healthier tea plants.

[Genetic diversity of Camellia sinensis germplasm in Guangdong Province based on morphological parameters and SRAP markers].

By the methods of phenotypic identification and SRAP makers amplification, the genetic diversity of twenty-five local tea cultivars in Guangdong Province and five contrastive cultivars from other regions was assessed and classified, and the phenotypic traits of the cultivars were clustered by Pearson correlation and Farthest neighbor methods. The coefficient of variation of the phenotypic traits was averagely 32.15%. Fine-hair had the highest coefficient of variation (42.41%), while the growth period of bud leaves had the smallest one (18.52%). Based on the cluster analysis of phenotypic traits, the test 30 tea cultivars could be clustered into 4 groups, 17 cultivars in the first group, 10 cultivars in the second group, 2 contrastive cultivars Yunnan-dayezhong and Lingyun-baimaocha in the third group, and 1 contrastive cultivar Hainan-dayezhong in the fourth group. After the amplification with 21 SRAP primers, a total of 127 fragments were detected, among which, 114 fragments were polymorphic, accounting for 88.67% of the total. The amplified fragments and polymorphic fragments per primer combination were averagely 6.05 and 5.43, respectively. At the genetic distance of 0.39 cm, the tea cultivars could be classified into three groups A, B and C, and 83.33% of the cultivars were belonged to group A. At the genetic distance of 0.31 cm, group A could be further classified into three sub-groups I , II and III, 13 cultivars in subgroup I, 2 cultivars in subgroup II, and 10 cultivars in subgroup III. It was not exactly the same between the clustering based on SRAP markers amplification and the performance of phenotypic traits.