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| 1 | The Reference Genome of Tea Plant and Resequencing of 81 Diverse Accessions Provide Insights into Its Genome Evolution and Adaptation显示文摘Tea plant is an important economic crop,which is used to produce the world's oldest and most widely consumed tea beverages.Here,we present a high-quality reference genome assembly of the tea plant(Camellia sinensis var.sinensis)consisting of 15 pseudo-chromosomes.LTR retrotransposons(LTR-RTs)account for 70.38%of the genome,and we present evidence that LTR-RTS play critical roles in genome size expansion and the transcriptional diversification of tea plant genes through preferential insertion in promoter regions and introns.Genes,particularly those coding for terpene biosynthesis pro-teins,associated with tea aroma and stress resistance were significantly amplified through recent tandem duplications and exist as gene clusters in tea plant genome.Phylogenetic analysis of the sequences of 81 tea plant accessions with diverse origins revealed three well-differentiated tea plant populations,support-ing the proposition for the southwest origin of the Chinese cultivated tea plant and its later spread to western Asia through introduction.Domestication and modern breeding left significant signatures on hundreds of genes in the tea plant genome,particularly those associated with tea quality and stress resis-tance.The genomic sequences of the reported reference and resequenced tea plant accessions provide valuable resources for future functional genomics study and molecular breeding of improved cul-tivars of tea plants. | Enhua Xia Wei Tong Yan Hou Yanlin An Linbo Chen Qiong Wu Yunlong Liu Jie Yu Fangdong Li Ruopei Li Penghui Li Huijuan Zhao Ruoheng Ge Jin Huang Ali Inayat Mallano Yanrui Zhang Shengrui Liu Weiwei Deng Chuankui Song Zhaoliang Zhang Jian Zhao Shu Wei Zhengzhu Zhang Tao Xia Chaoling Wei Xiaochun Wan | 2020 | Molecular Plant2020，13，7： | 51 |
| 2 | Attractive but Toxic: Emerging Roles of Glycosidically Bound Volatiles and Glycosyltransferases Involved in Their Formation显示文摘 | Chuankui Song Katja Hartl Kate McGraphery Thomas Hoffmann Wilfried Schwab | 2018 | Molecular Plant2018，11，10： | 16 |
| 3 | UGT74AF3 enzymes specifically catalyze the glucosylation of 4-hydroxy-2,5-dimethylfuran-3(2H)-one,an important volatile compound in Camellia sinensis显示文摘4-Hydroxy-2,5-dimethylfuran-3(2H)-one(HDMF)is an important odorant in some fruits,and is proposed to play a crucial role in the caramel-like notes of some teas.However,its biosynthesis and metabolism in tea plants are still unknown.Here,HDMF glucoside was unambiguously identified as a native metabolite in tea plants.A novel glucosyltransferase UGT74AF3a and its allelic protein UGT74AF3b specifically catalyzed the glucosylation of HDMF and the commercially important structural homologues 2(or 5)-ethyl-4-hydroxy-5(or 2)-methylfuran-3(2H)-one(EHMF)and 4-hydroxy-5-methylfuran-3(2H)-one(HMF)to their correspondingβ-D-glucosides.Site-directed mutagenesis of UGT74AF3b to introduce a single A456V mutation resulted in improved HDMF and EHMF glucosylation activity and affected the sugar donor preference compared with that of the wild-type control enzyme.The accumulation of HDMF glucoside was consistent with the transcript levels of UGT74AF3 in different tea cultivars.In addition,transient UGT74AF3a overexpression in tobacco significantly increased the HDMF glucoside contents,and downregulation of UGT74AF3 transcripts in tea leaves significantly reduced the concentration of HDMF glucoside compared with the levels in the controls.The identification of HDMF glucoside in the tea plant and the discovery of a novel-specific UDP-glucose:HDMF glucosyltransferase in tea plants provide the foundation for improvement of tea flavor and the biotechnological production of HDMF glucoside. | Yongxian Chen Xiangyang Guo Ting Gao Na Zhang Xiaochun Wan Wilfried Schwab Chuankui Song | 2020 | Horticulture Research2020，7，1： | 6 |
| 4 | Cold Stress-induced Glucosyltransferase CsUGT78A15 is Involved in the Formation of Eugenol Glucoside in Camellia sinensis显示文摘Eugenol is a natural phenolic compound known for its health-promoting properties and its ability to add a floral scent to tea plants.Plant eugenol glycosides have been identified and shown to make important contributions to fruit floral quality.However,the details of their biosynthesis and metabolism in tea plants are still unknown.Here,eugenol glucoside was unambiguously identified as a native metabolite in the tea plant,and its biosynthesis was shown to be induced by low temperature treatment.Through the analysis of UGTs induced by low temperature,the glycosyltransferase CsUGT78A15 was identified in tea,and its encoded protein was shown to catalyze the glucosylation of eugenol.Vmax/Km ratios showed that eugenol was the most suitable substrate for CsUGT78A15.Sugar donor preference analysis showed that CsUGT78A15 had a higher selectivity for glucose,followed by galactose and glucuronic acid.The expression of CsUGT78A15was correlatedwith the accumulation of eugenol glucoside in different tissues and genotypes of tea.Down-regulation of CsUGT78A15 led to a decreased eugenol glucoside content under cold stress,indicating that CsUGT78A15 plays an important role in the biosynthesis of eugenol glucoside under cold stress.The identification of eugenol glucoside in the tea plant and the discovery of a cold stress-induced eugenol glucosyltransferase in tea provide the foundation for the improvement of tea flavor under cold stress and the biotechnological production of eugenol glucoside. | Mingyue Zhao Binbin Cai Jieyang Jin Na Zhang Tingting Jing Jingming Wang Yuting Pan Zixiang Zhou Yifan Zhao Yingying Feng Feng Yu Mengting Zhang Yating Li Zhonghua Liu Chuankui Song | 2020 | Horticultural Plant Journal2020，6，6： | 4 |
| 5 | The effects of circadian rhythm on catechin accumulation in tea leaves显示文摘As the major bitter-and astringent-tasting components in tea leaves,catechins play an essential role in determining flavor characteristics and health benefits of tea products.This study analyzed the circadian rhythm effects on the accumulation pattern of catechins in tender leaves of three tea cultivars,'Huangkui'(HK),'Longjing43'(LJ43)and'Fuding Dabaicha'(FD).There were two major factors involved in the catechin regulation in tea plants:genetic background of the cultivar-specific regulation,and temporal effects of the circadian rhythm.Etiolated leaves of'HK'contained lower levels of catechins than'LJ43'and'FD'.The composition of EGCG in total catechins was significantly higher in'LJ43'leaves(65%)when compared with that in'HK'and'FD'(45%).The accumulation of individual catechins increased during the daytime but decreased overnight,especially over the time period of 22:00−02:00.The correlation analysis between catechins and environmental factors indicated catechin accumulation in tea leaves potentially resulted from a combined regulation of light and temperature.These findings provide new insights into our understanding of circadian rhythm regulation on flavor-important secondary metabolites in tea leaves. | Xiaohui Wang Wenfeng Zhu Xin Cheng Zhiqiang Lu Xuyang Liu Xiaochun Wan Chuankui Song Linlin Liu | 2021 | Beverage Plant Research2021，1，1： | 2 |
| 6 | Induction of priming by cold stress via inducible volatile cues in neighboring tea plants^FA显示文摘Plants have evolved sophisticated defense mechanisms to overcome their sessile nature.However,if and how volatiles from cold‐stressed plants can trigger interplant communication is still unknown.Here,we provide the first evidence for interplant communication via inducible volatiles in cold stress.The volatiles,including nerolidol,geraniol,linalool,and methyl salicylate,emitted from cold‐stressed tea plants play key role(s)in priming cold tolerance of their neighbors via a C‐repeatbinding factors‐dependent pathway.The knowledge will help us to understand how plants respond to volatile cues in cold stress and agricultural ecosystems. | Mingyue Zhao Lu Wang Jingming Wang Jieyang Jin Na Zhang Lei Lei Ting Gao Tingting Jing Shangrui Zhang Yi Wu Bin Wu Yunqing Hu Xiaochun Wan Wilfried Schwab Chuankui Song | 2020 | Journal of Integrative Plant Biology2020，62，10： | 2 |
| 7 | Promoter and coding sequence diversity of CsCCD1 may contribute to the differential accumulation of floralβ-ionone in fresh tea leaves显示文摘The carotenoid-derived volatileβ-ionone makes an important contribution to tea fragrance.Here,we qualitatively and quantitatively analysed 15 carotenoids in tea leaves of 13 cultivars by UHPLC-APCI-MS/MS.The 13 cultivars were divided into two groups by PCA(Principal Component Analysis)clustering analysis of their carotenoid content,and OPLS-DA(Orthogonal projections to latent structures)indicated that the levels ofβ-carotene(VIP=2.89)and lutein(VIP=2.30)were responsible for much of the variation between the two groups.Interestingly,theβ-carotene toβ-ionone conversion rates in Group 1 were higher than in Group 2,while theβ-carotene content was significantly lower in Group 1 than in Group 2.Theβ-ionone content was significantly higher in Group 1.Pearson Correlation Coefficient calculation between the transcription level of candidate genes(CsCCD1 and CsCCD4)and the accumulation ofβ-ionone indicated that CsCCD1 may involve in the formation ofβ-ionone in 13 cultivars.Prokaryotic expression and in vitro enzyme activity assays showed that‘Chuanhuang 1’had an amino acid mutation in carotenoid cleavage dioxygenases 1(CsCCD1)compared with‘Shuchazao’,resulting in a significantly higherβ-ionone content in‘Chuanhuang 1’.Sequence analysis showed that‘Chuanhuang 1’and‘Huangdan’had different CsCCD1 promoter sequences,leading to significantly higher CsCCD1 expression andβ-ionone accumulation in‘Chuanhuang 1’.These results indicated that the promoter and coding sequence diversity of CsCCD1 might contribute to the differential accumulation ofβ-ionone in different tea cultivars. | Jingming Wang Mingyue Zhao Ting Gao Yingying Feng Feiquan Wang Yuting Pan Jieyang Jin Tingting Jing Mengqian Lu Mengting Zhang Danyang Guo Xiaochun Wan Wilfried Schwab Chuankui Song | 2023 | Horticultural Plant Journal2023，9，4： | 1 |
| 8 | Amplification of early drought responses caused by volatile cues emitted from neighboring tea plants显示文摘Plants have developed sophisticated mechanisms to survive in dynamic environments.Plants can communicate via volatile organic compounds(VOCs)to warn neighboring plants of threats.In most cases,VOCs act as positive regulators of plant defense.However,the communication and role of volatiles in response to drought stress are poorly understood.Here,we showed that tea plants release numerous VOCs.Among them,methyl salicylate(MeSA),benzyl alcohol,and phenethyl alcohol markedly increased under drought stress.Interestingly,further experiments revealed that drought-induced MeSA lowered the abscisic acid(ABA)content in neighboring plants by reducing 9-cis-epoxycarotenoid dioxygenase(NCED)gene expression,resulting in inhibition of stomatal closure and ultimately decreasing early drought tolerance in neighboring plants.Exogenous application of ABA reduced the wilting of tea plants caused by MeSA exposure.Exposure of Nicotiana benthamiana to MeSA also led to severe wilting,indicating that the ability of drought-induced MeSA to reduce early drought tolerance in neighboring plants may be conserved in other plant species.Taken together,these results provide evidence that drought-induced volatiles can reduce early drought tolerance in neighboring plants and lay a novel theoretical foundation for optimizing plant density and spacing. | Jieyang Jin Mingyue Zhao Ting Gao Tingting Jing Na Zhang Jingming Wang Xianchen Zhang Jin Huang Wilfried Schwab Chuankui Song | 2021 | Horticulture Research2021，8，1： | 1 |
| 9 | Correction: Amplification of early drought responses caused by volatile cues emitted from neighboring tea plants显示文摘After the publication of this article 1,the authors became aware that the word of“tea”was not mentioned in the title.To make the title clear and complete,the word‘tea’is added to the title. | Jieyang Jin Mingyue Zhao Ting Gao Tingting Jing Na Zhang Jingming Wang Xianchen Zhang Jin Huang Wilfried Schwab Chuankui Song | 2021 | Horticulture Research2021，8，1： | 1 |
| 10 | A geraniol synthase regulates plant defense via alternative splicing in tea plants显示文摘Geraniol is an important contributor to the pleasant floral scent of tea products and one of the most abundant aroma compounds in tea plants;however,its biosynthesis and physiological function in response to stress in tea plants remain unclear.The proteins encoded by the full-length terpene synthase(CsTPS1)and its alternative splicing isoform(CsTPS1-AS)could catalyze the formation of geraniol when GPP was used as a substrate in vitro,whereas the expression of CsTPS1-AS was only significantly induced by Colletotrichum gloeosporioides and Neopestalotiopsis sp.infection.Silencing of CsTPS1 and CsTPS1-AS resulted in a significant decrease of geraniol content in tea plants.The geraniol content and disease resistance of tea plants were compared when CsTPS1 and CsTPS1-AS were silenced.Down-regulation of the expression of CsTPS1-AS reduced the accumulation of geraniol,and the silenced tea plants exhibited greater susceptibility to pathogen infection than control plants.However,there was no significant difference observed in the geraniol content and pathogen resistance between CsTPS1-silenced plants and control plants in the tea plants infected with two pathogens.Further analysis showed that silencing of CsTPS1-AS led to a decrease in the expression of the defense-related genes PR1 and PR2 and SA pathway-related genes in tea plants,which increased the susceptibility of tea plants to pathogens infections.Both in vitro and in vivo results indicated that CsTPS1 is involved in the regulation of geraniol formation and plant defense via alternative splicing in tea plants.The results of this study provide new insights into geraniol biosynthesis and highlight the role of monoterpene synthases in modulating plant disease resistance via alternative splicing. | Hao Jiang Mengting Zhang Feng Yu Xuehui Li Jieyang Jin Youjia Zhou Qiang Wang Tingting Jing Xiaochun Wan Wilfried Schwab Chuankui Song | 2023 | Horticulture Research2023，10，10： | 0 |
| 11 | Molecular mechanism of lycopene cyclases regulating carotenoids ratio in different branches during tea leaf and flower development显示文摘Carotenoids are essential components in tea quality, contributing to leaf color and aroma. However, little information about carotenoids in different tea cultivars and their biosynthesis regulation mechanism during leaf development is known. Here we analyzed carotenoids by HPLC in the buds and leaves of 113 tea cultivars harvested on the same day. By profile clustering, carotenoids were divided into five groups. Same group cultivars displayed divergence in the total content of carotenoids but a similar molar ratio. To figure out the molecular mechanisms of this phenomenon, we further characterized all functional lycopene cyclases, which are the branch point of the carotenoid biosynthesis pathway. Two β-lycopene cyclases(CsLCYB1 and CsLCYB2) and one ε-lycopene cyclase(CsLCYE1) were cloned. Subcellular localization analysis showed that all cloned CsLCYs were localized in plastids. Enzyme activity assays in E. coli indicated both CsLCYBs catalyzed lycopene into β-carotene, and CsLCYE1 produced δ-carotene and ε-carotene. We found CsLCYB1 and CsLCYE1 predominantly expressed in leaf, while CsLCYB2was mainly expressed during flowering stages. Suppression by antisense oligonucleotides reduced CsLCYB1 and CsLCYE1 transcripts and led to reduction of both β,β-branch and β,ε-branch carotenoids in leaf. The expression levels of CsLCYB1 showed a significant positive correlation withβ,β-branch carotenoids in leaf. Our study provides carotenoid profiles of different tea cultivars, which can assist tea producers in selecting cultivars of interest. Meanwhile, we proposed the molecular mechanism of carotenoids reflecting the tenderness of tea plant leaf from a metabolic flux perspective, and suggested lycopene cyclase that could be applied to the breeding of tea varieties with different branch carotenoids. | Anqi Peng Xiaoyan Tang Yingying Feng Yun Huang Jilai Cui Kai Tian Mengqian Lu Yifan Zhao Yuting Pan Sanjin Wang Yutong Hu Wei Zhao Chuankui Song Qiang Wang | 2023 | Horticultural Plant Journal2023，9，6： | 0 |
| 12 | Volatile compound-mediated plant-plant interactions under stress with the tea plant as a model显示文摘Plants respond to environmental stimuli via the release of volatile organic compounds(VOCs),and neighboring plants constantly monitor and respond to these VOCs with great sensitivity and discrimination.This sensing can trigger increased plant fitness and reduce future plant damage through the priming of their own defenses.The defense mechanism in neighboring plants can either be induced by activation of the regulatory or transcriptional machinery,or it can be delayed by the absorption and storage of VOCs for the generation of an appropriate response later.Despite much research,many key questions remain on the role of VOCs in interplant communication and plant fitness.Here we review recent research on the VOCs induced by biotic(i.e.insects and pathogens)and abiotic(i.e.cold,drought,and salt)stresses,and elucidate the biosynthesis of stress-induced VOCs in tea plants.Our focus is on the role of stress-induced VOCs in complex ecological environments.Particularly,the roles of VOCs under abiotic stress are highlighted.Finally,we discuss pertinent questions and future research directions for advancing our understanding of plant interactions via VOCs. | Jieyang Jin Mingyue Zhao Tingting Jing Mengting Zhang Mengqian Lu Guomeng Yu Jingming Wang Danyang Guo Yuting Pan Timothy D.Hoffmann Wilfried Schwab Chuankui Song | 2023 | Horticulture Research2023，10，9： | 0 |
| 13 | Subfunctionalization of a monolignol to a phytoalexin glucosyltransferase is accompanied by substrate inhibition显示文摘Uridine diphosphate-dependent glycosyltransferases(UGTs)mediate the glycosylation of plant metabolites,thereby altering their physicochemical properties and bioactivities.Plants possess numerous UGT genes,with the encoded enzymes often glycosylating multiple substrates and some exhibiting substrate inhibition kinetics,but the biological function and molecular basis of these phenomena are not fully understood.The promiscuous monolignol/phytoalexin glycosyltransferase NbUGT72AY1 exhibits substrate inhibition(Ki)at 4 mM scopoletin,whereas the highly homologous monolignol StUGT72AY2 is inhibited at 190 mM.We therefore used hydrogen/deuterium exchange mass spectrometry and structure-based mutational analyses of both proteins and introduced NbUGT72AY1 residues into StUGT72AY2 and vice versa to study promiscuity and substrate inhibition of UGTs.A single F87I and chimeric mutant of NbUGT72AY1 showed significantly reducedscopoletin substrate inhibition,whereas its monolignolgly cosylation activity was almost unaffected.Reverse mutations in StUGT72AY2 resulted in increased scopoletin glycosylation,leading to enhanced promiscuity,which was accompanied by substrate inhibition.Studies of 3D structures identified open and closed UGT conformers,allowing visualization of the dynamics of conformational changes that occur during catalysis.Previously postulated substrate access tunnels likely serve as drainage channels.The results suggest a two-site model in which the second substrate molecule binds near the catalytic site and blocks product release.Mutational studies showed that minor changes in amino acid sequence can enhance the promiscuity of the enzyme and add new capabilities such as substrate inhibition without affecting existing functions.The proposed subfunctionalization mechanism of expanded promiscuity may play a role in enzyme evolution and highlights the importance of promiscuous enzymes in providing new functions. | Jieren Liao Guangxin Sun Elisabeth Kurze Wieland Steinchen Timothy D.Hoffmann Chuankui Song Zhiwei Zou Thomas Hoffmann Wilfried G.Schwab | 2023 | Plant Communications2023，4，3： | 0 |
| 14 | First report of Muyocopron laterale causing a new leaf disease of Camellia sinensis in China显示文摘Tea plant(Camellia sinensis)diseases are one of the factors that reduce tea production yield and quality.Herein,a new leaf disease of tea plants was observed in tea plantations.A representative isolate was obtained from diseased leaf by the traditional fungus separation method.The isolate identified was confirmed as Muyocopron laterale based on morphological and molecular results and phylogenetic tree analysis.Pathogenicity tests were conducted on tea plant seedlings,and which is fulfilling Koch’s postulates.The disease was first identified in C.sinensis leaves caused by M.laterale in China.In the future,the results of this studies will enrich our knowledge and help control tea leaf disease. | Hao Jiang Mengting Zhang Youjia Zhou Xuehui Li Jiaxun Li Chuankui Song | 2022 | Beverage Plant Research2022，2，1： | 0 |
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