Journal of Tea Science

Identification of Pectin Methylesterase and Its Inhibitory Subfamily Genes, and Functional Analysis of CsPME55 in Response to Fluoride Stress in Camellia sinensis

XU Wenluan, WEN Xiaoju, JIA Yuxuan, NI Dejiang, WANG Mingle, CHEN Yuqiong

Journal of Tea Science. 2024, 44(6): 869-886. doi:10.13305/j.cnki.jts.2024.06.008

Abstract ( 56 )

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Tea plant (Camellia sinensis) has fluoride (F) enrichment characteristics, and F is mainly enriched in the cell wall component pectin. Pectin methylsterase (PME) and its inhibitor PMEI can catalyze the modification of pectin, thereby affecting cell wall characteristics and participating in the regulation of processes like plant growth and development, stress response and so on. In this study, 85 CsPMEs and 56 CsPMEIs were identified from the C. sinensis ‘Shuchazao’ genome, which were divided into 4 and 5 subgroups, respectively. Distinct subgroups may exhibit functional distinction due to varied gene architectures, conserved motifs and expression patterns. Quantitative real-time PCR (qRT-PCR) analysis reveals that the expression levels of CsPME3a, CsPME55, CsPMEI1 and CsPMEI3 were significantly induced in the mature leaves of ‘Fuding Dabaicha’ under F treatment. Moreover, overexpression of CsPME55 alleviated Arabidopsis root growth inhibition induced by F stress, suggesting its potential role in F stress regulation in tea plants. These findings could pave the way for further research on the functional involvement of PME and PMEI gene families in F response.

Studies on the Regulation of EGCG Biosynthesis in Tea Plants by Potassium Nutrition

YANG Nan, LI Zhuan, LIU Meichen, MA Junjie, SHI Yuntao, WEI Xiangning, LIN Yangshun, MAO Yuyuan, GAO Shuilian

Journal of Tea Science. 2024, 44(6): 887-900. doi:10.13305/j.cnki.jts.2024.06.012

Abstract ( 56 )

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Epigallocatechin gallate (EGCG) is an important flavor and health functional component in tea. Previous studies have found that EGCG biosynthesis in tea plants is affected by potassium nutrition, but the regulatory mechanism of its biosynthesis is currently unclear. This study used one year old tea seedlings of Huangdan as the experimental object, and set up 5 treatment groups (K1-K5), with K₂SO₄ concentrations of 0.4, 0.6, 0.8, 1.0 mmol∙L^-1 and 1.2 mmol∙L^-1 for irrigation, respectively. The joint analysis of transcriptomics and metabolomics shows that, under low-potassium treatment (K1), the flavonoid contents in the new shoots of tea plants accumulated significantly and the EGCG content reached the highest level, and the difference reached a significant level compared with that of the high potassium treatment (K5). The related metabolites of phenylalanine, cinnamic acid and p-coumaric acid on the EGCG synthesis pathway were up-regulated in the K4 or K5 treatments, whereas the downstream metabolites of the flavonoid pathway (dihydroquercetin, dihydromyricetin, colorless delphinidin pigment and epigallocatechin) were up-regulated in the K1 and K2 treatments. Under the influence of potassium nutrition, EGCG biosynthesis was positively regulated by a series of structural genes CsCHI, F3′5′H, CsF3H (CSS0019002), CsANS, CsANR, Csaro DE, CsSCPL, and transcription factor (MYB306), as well as negatively regulated by CsPAL, CsC4H, Cs4CL, CsCHS, CsF3H (CSS0016177), CsDFR (CSS0011557) and transcription factor (NAC83). It is thus clear that potassium nutrition regulates EGCG synthesis by affecting the expressions of key genes in tea plants, thereby affecting EGCG content. This study provided a scientific basis for the regulation of EGCG biosynthesis in tea plants by potassium nutrition.

Effects of Combined Drought and Low-temperature Stress on Photosynthetic Physiological Characteristics of Tea Plants and Simulation Prediction

ZHAO Qian, LIU Qian, CAI-HE Jiayi, HE Jieqi, FANG Yunya, LIU Yuxin, CHEN Chao, ZHENG Yaodong, ZHANG Tianjing, YU Wenjuan, YANG Guang

Journal of Tea Science. 2024, 44(6): 901-916. doi:10.13305/j.cnki.jts.2024.06.010

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This study aimed to investigate the effects of multiple climatic stresses on the photosynthetic efficiency of tea plants and to devise an efficient, precise stress classification system for real-time monitoring. We focused on the typical tea cultivars grown extensively in Fujian Province and systematically monitored their photosynthetic physiological responses under combined drought and low-temperature stress. Utilizing the collected data, we established a rapid stress classification method based on photosynthetic physiological characteristics and constructed a photosynthesis prediction and early warning model. The results reveal that all tested tea cultivars exhibited a significant decline in leaf photosynthetic efficiency under combined stress, with the decreasing trend displaying a clear linear relationship with increasing stress intensity. Notably, ‘Tieguanyin’ demonstrated a significantly lesser decline in photosynthetic efficiency compared to other cultivars, suggesting its robust stress tolerance. In contrast, ‘Fuding Dabaicha’ showed the least stress tolerance. By selecting photosynthetic physiological parameters highly sensitive to combined stress and employing the K-means clustering algorithm, we developed a rapid stress classification method with an accuracy exceeding 80%. Various models were then used to predict and validate the response of photosynthetic physiological indicators to environmental stress, with the Random Forest (RF) model yielding the highest accuracy. This study provided a reference for the selection and breeding of tea cultivars under diverse climatic conditions. The stress classification method enables swift categorization of combined stress in tea plants, while the RF model facilitates non-destructive monitoring and early warning of photosynthetic physiology, offering significant practical value in tea production.

Changes in Cell Wall Structure and Photosynthetic Characteristics of Tea Leaves under Low Temperature Stress

LIU Xiaolu, ZHU Yalan, YU Min, GAI Xinyue, FAN Yangen, SUN Ping, HUANG Xiaoqin

Journal of Tea Science. 2024, 44(6): 917-927. doi:10.13305/j.cnki.jts.2024.06.001

Abstract ( 56 )

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To investigate the molecular mechanisms of tea plants in response to low temperature stress, this study simulated the spring chill temperature pattern, using ‘Fuding Dabaicha’ as the experimental material for transcriptome sequencing, and subjected it to varying low temperature treatments. Differentially expressed genes (DEGs) were analyzed using the GO and KEGG pathway databases for metabolic pathway enrichment analysis, which revealed that these genes were mainly enriched in plant cell wall and metabolic pathways related to photosynthesis. Subsequently, thirteen DEGs were selected for validation via real-time quantitative PCR (qPCR), confirming the consistency of the qPCR results with the transcriptome sequencing data, thereby validating the reliability of the transcriptome data. In a subsequent study, two tea cultivars, ‘Fuding Dabaicha’ and ‘Shuchazao’, were used to evaluate various physiological indices, including leaf tissue structure, the contents of various cell wall components (cellulose, hemicellulose, and pectin), chlorophyll content, and chlorophyll fluorescence parameters. The results indicate that the leaf tissue structures of both tea cultivars underwent different degrees of thickening in response to low-temperature stress. Notably, significant differences were observed in the contents of cellulose and hemicellulose between the two cultivars, whereas the pectin content change was less pronounced. Furthermore, the chlorophyll content, photochemical quenching coefficient, maximum photochemical efficiency, and relative electron transport rate all exhibited a downward trend. Conversely, the non-photochemical quenching coefficient showed an upward trend. These observations highlight the key role of changes in cell wall components, particularly hemicellulose, and changes in photosynthesis-related parameters in the tea plants’ response to low temperature.

Effect of Bamboo Density on the Physiological Growth and Tea Quality of Tea Plants under the Moso Bamboo Forest

ZHAO Jiancheng, NI Huijing, WANG Bo, CAI Chunju, YANG Zhenya

Journal of Tea Science. 2024, 44(6): 928-940. doi:10.13305/j.cnki.jts.2024.06.003

Abstract ( 43 )

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To explore the adaptive mechanism of physiological growth of tea plants under different bamboo density conditions, three treatments, namely T1, T2 and T3, were set in the natural mixed forest of moso bamboo (Phyllostachys edulis) and tea plants [Camellia sinensis (L.) O. Kuntze] in the Mogan mountain area of Zhejiang Province, with bamboo densities of 1 200 plants·hm^-2, 1 800 plants·hm^-2 and 2 400 plants·hm^-2, respectively, and 3 000~3 300 plants·hm^-2 as the control (CK). Tea yield was evaluated, nitrogen, phosphorus, nonstructural carbohydrates (NSC) and biomass distribution of tea tree were determined, root morphology, distribution and architecture were analyzed, and quality indicators of tea such as amino acids, theanine and tea polyphenols were tested after 2 years of cultivation. The results show that (1) compared to CK, T2 and T3 treatments significantly increased tea yield by 15.96% and 18.8%, respectively, increased biomass of leaves and lateral roots, and increased the nitrogen, phosphorus and NSC contents in leaves, (2) T1, T2 and T3 treatments significantly increased the root length, root surface area and root length ratio of the >0-1 mm diameter grade in 0-20 cm soil layer, reduced root diameter and root length ratio of thick roots (diameter>2 mm) in 0-20 cm soil layer, increased the contents of starch and soluble sugars in lateral roots, reduced the topological index of the whole root system and increased the root tips and root fractal dimension, (3) T1 and T2 treatments significantly increased the contents of amino acids, caffeine and tea polyphenols and water extract, and reduced the polyphenols-amino acid ratio. In conclusion, reasonable control of bamboo density can promote the growth of tea plants and improve the tea yield effectively. Tea plants promote the growth of leaves and lateral roots by increasing nitrogen, phosphorus and non-structural carbohydrate accumulation in leaves and lateral roots and promote root encroachment of surface soil space by increasing the proportion of fine roots, enhancing the root branch strength, reducing the cost of root construction to accommodate moderate closure conditions and interspecific competition strength. The conclusion can provide scientific basis for the management of innovation mode of tea plants under the moso bamboo forest.

The Orientation and Landing of the Spiny Citrus Whiteflies Attracted by Tea Cultivar ‘Dahuangpao’

WU Yiqi, CHENG Yanjun, LIANG Yueer, LU Jianliang, HAN Shanjie, HAN Baoyu

Journal of Tea Science. 2024, 44(6): 941-948. doi:10.13305/j.cnki.jts.2024.06.002

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‘Dahuangpao’ is a yellowing new tea elite cultivar with special yellow foliage. The adults of spiny citrus whitefly, Aleurocanthus spiniferus, is keen on flying to tea cultivars whose leaf color is bright and shining with yellow chroma being slightly high. Especially, the whiteflies preferred to directionally fly to and gather on ‘Dahuangpao’ tea shoots for mating, chasing, oviposition, piercing and sucking sap, secreting and resting. In order to reveal its taxis mechanism, the volatiles of the tea seedlings were collected by headspace aeration sampling method from which twenty compounds were identified. Among them methyl salicylate, linalool, cis-3-hexen-1-ol, n-hexanol and tran-2-hexenal significantly attracted the adults of the whiteflies in a Y-tube olfactometer bioassay, respectively. Within resting habitat in greenhouse, ‘Dahuangpao’ tea seedling models baited by the five infochemicals at equal ratio intensely attracted the adults in flight. This study demonstrated the five volatile compounds from ‘Dahuangpao’ tea seedlings were strongly attractive to the adults. And the combination of these volatiles with the special yellow leaf colour, and the shape of tea plants resulted in orientation and landing of adults onto ‘Dahuangpao’ tea plants.

Research on Tea Bud Recognition Based on Improved YOLOv8n

YANG Xiaowei, SHEN Qiang, LUO Jinlong, ZHANG Tuo, YANG Ting, DAI Yuqiao, LIU Zhongying, LI Qin, WANG Jialun

Journal of Tea Science. 2024, 44(6): 949-959. doi:10.13305/j.cnki.jts.2024.06.005

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Accurate recognition of tea buds in complex natural environment is one of the key technologies to realize intelligent picking of tea buds by agricultural robots. To address the problem of low recognition accuracy of tea buds in complex environment of tea gardens, a tea bud recognition method based on improved YOLOv8n was proposed. The Honor Mobile Phone was used to collect the RGB images of tea buds, and the image annotation of tea buds was completed. The labeled data was divided according to the 8∶1∶1 radio of the training set and test set. To effectively extract bud features and reduce model redundancy calculation and memory access, FasterNet was used to replace the backbone network of YOLOv8n model for feature extraction. To suppress the background information of the tea garden environment and enhance the feature extraction ability of tea buds, the global attention mechanism (GAM) module was introduced at the end of the backbone network (after the SPPF module). To further improve the recognition accuracy of tea buds, the Context Guided (CG) module was introduced into the Neck network to learn the joint features of local features and surrounding environment of tea buds. The improved YOLOV8n algorithm was trained and tested by using the constructed tea bud data set. The ablation experiments verify that the FasterNet network, GAM attention mechanism and CG module effectively improved the recognition accuracy of the YOLOv8n model. The mean average accuracy (mAP) of the improved YOLOv8n model on the multi-category tea bud data set was 94.3%. Compared with the original YOLOv8n model, the mAP of single bud, one bud and one leaf, and one bud and two leaves of tea buds increased by 2.2, 1.6 and 2.7 percentage points, respectively. The improved YOLOv8n model was tested for performance comparison with YOLOv3-tiny, YOLOv3, YOLOv5m, YOLOv7-tiny, YOLOv7 and YOLOv8n models. The experimental results show that the improved YOLOv8n model has a higher accuracy in identifying tea buds. The experimental results demonstrate that the improved YOLOv8n model can effectively improve the accuracy of tea bud recognition and provide technical support for intelligent tea picking robots.

Effects of Different Charcoal Baking Times on the Sensory Quality and Volatile Compounds of Dahongpao

XIE He, XIAO Han, HU Tengfei, CHEN Guohe, LIU Yang, OU Xingchang, JIANG Ronggang, YU Liming, LI Qin, HUANG Jian'an, LIU Zhonghua, WANG Chao

Journal of Tea Science. 2024, 44(6): 960-972. doi:10.13305/j.cnki.jts.2024.06.011

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To investigate the aroma characteristics and dynamic changes of volatile compounds of Dahongpao at different charcoal baking times, quantitative descriptive analysis (QDA), headspace solid-phase microextraction (HS-SPME) and comprehensive two-dimensional gas chromatography-quadrupole time-of-flight mass spectrometry (GC×GC-Q-TOF/MS) were applied to analyze the volatile compounds at different charcoal baking times of 200 min, 300 min, 400 min, 500 min and 600 min (LF200, LF300, LF400, LF500 and LF600). The results show that the grassy aroma appeared in the early stage of charcoal baking, and with the extension of charcoal baking time, the grassy aroma gradually disappeared, and the floral and fruity aroma were prominent, and the floral and fruity aroma were the most porminent in the LF600 sample. A total of 304 volatile compounds were detected in Dahongpao at different charcoal baking times, among which alcohols, aldehydes and esters were the main ones. The principal component analysis (PCA) and hierarchical cluster analysis (HCA) results divided the samples under different carbon baking times into three groups, and 11 major differential compounds were screened by variable important for the projection (VIP)>1 and relative odor activity value (ROAV)>1. Among them, the relative content of (E,E)-2,4-hexadienal in the grassy characteristic decreased with the extension of the charcoal baking time, while the relative contents of methyl benzoate, methyl phenylacetate, nerolidol and myrcene in the fruity characteristics increased with the extension of the charcoal baking time. The purpose of this study was to regulate the quality of Dahongpao, and to improve the drinking attributes for the subsequent production and processing of Dahongpao.

Optimization and Application of Analysis Method for Volatile Enantiomers in ‘Jinxuan’ Roasted Green Tea

ZHANG Shuyi, MA Chengying, CHEN Wei, MIAO Aiqing, QIAO Xiaoyan, LIN Dongchun, XIA Hongling, XU Jingyi

Journal of Tea Science. 2024, 44(6): 973-984. doi:10.13305/j.cnki.jts.2024.06.006

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To investigate the changes in volatile enantiomers during the drying of ‘Jinxuan’ roasted green tea, an optimized method using headspace solid-phase microextraction-enantioselective-gas chromatography-mass spectrometry (HS-SPME-Es-GC-MS) was applied for qualitative and quantitative enantiomer analysis. The chemometric evaluation was also performed on four roasted green tea samples dried at 120 ℃ for different durations (30 min, 60 min, 90 min and 120 min). The optimal HS-SPME conditions were found to be: addition of NaCl (3 mL, 3 mol·L^-1, solid-to-liquid ratio of 1∶6, extraction temperature of 60 ℃, and extraction time of 25 min. The optimal temperature program for Es-GC-MS was as follows: initial temperature at 35 ℃ held for 2 min, then ramped up to 110 ℃ at 4 ℃ min^-1 and held for 10 min, followed by a further increase to 210 ℃ at 4 ℃ min^-1. Using an optimized method to analyze four ‘Jinxuan’ roasted green tea samples, a total of 8 enantiomers were detected, with no change in the types as drying time increased. Among them, the contents of R-(-)-linalool and S-(+)-linalool increased with the drying process, while the contents of the other 6 enantiomers began to decline after 60 min or 90 min. Hierarchical cluster analysis and partial least squares discrimination analysis show that the samples with different drying times were obviously classified into four groups. Based on variable importance in projection (VIP) >1 and significance analysis, four different enantiomers were screened, and their VIP values were ranked in descending order as R-(-)-linalool, (2R,5S)-theaspirane B, (2R,5R)-theaspirane A, and S-(+)-linalool. This study provided a reference for improving aroma quality and selecting drying processes during the roasting of green tea.

Evaluation of Aroma Types and Key Aroma Components of Bud-leaf Type Yellow Tea Based on GC-O-MS

ZHOU Hongyu, WANG Yuanyuan, LI Faxin, NIE Congning, FENG Dejian, ZHAO Yueling, DU Xiao

Journal of Tea Science. 2024, 44(6): 985-1004. doi:10.13305/j.cnki.jts.2024.06.004

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To comprehensively analyze the characteristic aroma components of bud-leaf type yellow tea, headspace solid-phase microextraction (HS-SPME) combined with gas chromatography-mass spectrometry (GC-MS), gas chromatography-olfactometry (GC-O), sensory evaluation and chemometrics were used to determine the aroma components of bud-leaf type yellow tea. It was found that the bud-leaf yellow tea was mainly characterized by a clear and sweet aroma, which was divided into three categories of “fresh-tender aroma”, “chestnut-sweet aroma” and “sweet aroma” by sensory evaluation. A total of 119 volatile components were detected by GC-MS, and then the candidate volatiles were screened out by the VIP values of cluster analysis (HCA) and orthogonal partial least squares discriminant analysis (OPLS-DA). Further, ROAV with GC-O analysis was used to clarify the key aroma components of bud-leaf type yellow tea. There were 60 characteristic aroma components with ROAV>1 in the bud-leaf type yellow tea, among which aldehydes dominated the “fresh-tender” type, alcohols dominated the “chestnut-sweet” type, 2-ethyl-5-methylpyrazine and 2,3-diethyl-5-methylpyrazine contributed significantly to the “sweet” type. The olfactory validation indicates that phenylethanol was the most important component for the fresh and sweet aroma of yellow tea. The key aroma of “fresh-tender” included phenylethanol, jasmine ketone, styrene, linalool, nonanal. The “chestnut-sweet aroma” was mainly derived from the components of heptanal, β-damascenone and 2-n-pentylfuran, while the “sweet aroma” was dominated by pyrazines and aldehydes, which presented a unique sweet and caramelized aroma. This study reveals the diversity of aroma components in bud-leaf type yellow tea, and elucidates the aroma differences between aroma types and their causes.

Neuroprotective Mechanisms of Aged Liupao Tea against Aβ_25-35-induced PC12 Cell Damage

NIE Qing, PANG Yuelan, WU Huan, DING Shuqia, ZHONG Keyu, LIU Zhonghua, CAI Shuxian

Journal of Tea Science. 2024, 44(6): 1005-1013. doi:10.13305/j.cnki.jts.20241203.001

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In this study, an Aβ_25-35-induced PC12 cell damage model was established to investigate the neuroprotective effects and underlying mechanisms of aged Liupao tea (ALPT), with green tea (GT) as a reference. The results show that Aβ_25-35 significantly reduced PC12 cell viability, induced mitochondrial dysfunction, and promoted the formation of toxic aggregates and related pathways. ALPT markedly improved cell survival, increased mitochondrial membrane potential, and significantly inhibited the accumulation of toxic aggregates and the formation of related pathways. Furthermore, transcriptome analysis reveals that the overall gene expression pattern in the ALPT treatment group was the opposite to that in the Aβ_25-35 group, with upregulated genes involved in mitophagy, glycolysis and glycerophospholipid metabolism, and downregulated genes associated with cell cycle regulation, ribosomal function, ubiquitin-mediated proteolysis and cellular senescence. Overall, both GT and ALPT exhibited significant protective effects against Aβ_25-35-induced PC12 cell damage, though transcriptomic differences suggest that ALPT may have superior bioavailability due to its active components. This study provided experimental evidence for the potential application of ALPT in the prevention and treatment of neurodegenerative diseases.

Determination of Selenium in Tea by Microwave Digestion-inductively Coupled Plasma Mass Spectrometry (ICP-MS)

XU Wenqi

Journal of Tea Science. 2024, 44(6): 1014-1022. doi:10.13305/j.cnki.jts.2024.06.007

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In order to improve the sensitivity and stability of microwave digestion-inductively coupled plasma mass spectrometry (ICP-MS), an accurate method for the determination of selenium in tea was established. The sensitization effects of methanol, ethanol, isopropanol, n-butanol etc. as sensitizers were investigated, as well as the impact of adding 0-10% n-butanol to the internal standard on the accurate determination of total selenium content in tea were tested. By optimizing the pretreatment method and ICP-MS instrument parameters, a method for the accurate detection of total selenium content in tea using microwave digestion-ICP-MS with 3% n-butanol as the sensitizer was established. This method enhances the selenium signal intensity, thereby improving the accuracy and stability of the total selenium content detection in tea. To determine the optimum conditions for the determination of total selenium in tea, n-Butanol as sensitizer had lower detection limit and higher accuracy (0.001 71 mg·kg^-1), and 3% n-butanol had the best sensitizing effect. It was more accurate to determine the selenium content in tea reference materials. The linear correlation coefficient R² of selenium curve was greater than 0.999. The recovery of the added standard was from 86.0% to 104%, and the relative standard deviations (RSD) was between 1.10% and 4.44%. The paired t-test shows that the P value was greater than 0.05, and there was no significant difference between the two atomic fluorescence spectroscopy. This method is simple, showing high sensitivity, low detection limit, high precision and good reproducibility. This method is suitable for the accurate determination of selenium in tea.

An Empirical Study on the Increase in Purchase Intention of Novel-tea Beverage among Young Consumers: Based on the Perspective of Customer Perceived Value

YAN Pengxiang, PENG Kang, CHEN Fuqiao, JIANG Renhua

Journal of Tea Science. 2024, 44(6): 1023-1036. doi:10.13305/j.cnki.jts.2024.06.009

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In recent years, the novel-tea beverage industry has developed rapidly, becoming the third largest form of tea consumption after traditional loose-leaf tea and fast-moving bottled tea. The rapid expansion has intensified competition within the industry, making it crucial to increase customers’ purchase intention and frequency for the long-term healthy development of the industry. This study constructed an analytical model for enhancing the purchase intention of young consumers of novel-tea beverage from the perspective of customer perceived value. Though empirical research based on 1 900 consumer survey data points, the study examined the impact of product innovation, scene experience and brand exposure on customers’ purchase intention. The findings are as follows: (1) Product innovation, scene experience and brand exposure all positively influence customers’ purchase intention. (2) Functional value, experiential value and emotional value enhance the effects of product innovation, scene experience and brand exposure on purchase intention. (3) Product innovation, scene experience and brand exposure all positively affect purchase frequency, though there is some discerpancy between purchase intention and actual purchase behavior. Notably, product innovation plays a significant role in translating purchase intention into actual purchase behavior. Based on these conclusions, it is suggested that novel-tea beverage companies should highlight the uniqueness of product innovation, build interesting scene experiences, and link brand exposure emotions in their operations.

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