Ahammed G J, Wu Y X, Wang Y M, Guo T M, Shamsy R, Li X. 2023. Epigallocatechin-3-gallate (EGCG): A unique secondary metabolite with diverse roles in plant–environment interaction. Environmental and Experimental Botany, 209, 105299.

Akanvou R, Bastiaans L, Kropff M J, Goudriaan J, Becker M. 2001. Characterization of growth, nitrogen accumulation and competitive ability of six tropical legumes for potential use in intercropping systems. Journal of Agronomy and Crop Science, 187, 111–120.

Azapagic A, Bore J, Cheserek B, Kamunya S, Elbehri A. 2016. The global warming potential of production and consumption of Kenyan tea. Journal of Cleaner Production, 112, 4031–4040.

Boonerjee S, Islam M A, Islam S M M, Paul S K, Uddin M T, Alamgir M S. 2025. Consumption and export potential of tea in bangladesh: A field study. Journal of Agriculture and Food Research, 19, 101607.

Brooker R W, Bennett A E, Cong W, Daniell T J, George T S, Hallett P D, Hawes C, Iannetta P P M, Jones H G, Karley A J, Li L, McKenzie B M, Pakeman R J, Paterson E, Schöb C, Shen J, Squire G, Watson C A, Zhang C, Zhang F, et al. 2015. Improving intercropping: A synthesis of research in agronomy, plant physiology and ecology. New Phytologist, 206, 107–117.

Cao R, Yang J C, Meng Z T, Zhou H, Dong X, Peng L, Hou S S. 2025. Linking microbial communities and organic matter dynamics in longjing and fuding tea ecosystems. Journal of Food Biochemistry, 2025, 9981444.

Chen D, Dou Q P. 2008. Tea polyphenols and their roles in cancer prevention and chemotherapy. International Journal of Molecular Sciences, 9, 1196–1206.

Chen M J, Zhou P Q, Bao Q, Wang H, Wang Y J, Fu H P. 2025. Intercropping different legumes in tea plantation improves soil properties and tea quality components by regulating rhizosphere soil microorganisms. Agronomy, 15, 511.

Chen Y, Li J W, Yang N, Hu Z H, Luo W, Chen C, Wang Y H, Chen X, Li X H, Zhuang J. 2025. Nitrogen forms and nitrogen deficiency regulate theanine accumulation patterns in tea plants (Camellia sinensis) during winter dormancy. Horticulturae, 11, 444.

Chen Y Y, Li J L, Zhou B, Wu X M, Cui Y Y, Feng S M, Hu H T, Tang J C. 2023. Effects of intercropping with Vulpia myuros in tea plantation on soil and tea quality components. Scientia Agricultura Sinica, 56, 4916–4929. (in Chinese)

Cong W F, Hoffland E, Li L, Janssen B H, van der Werf W. 2015. Intercropping affects the rate of decomposition of soil organic matter and root litter. Plant and Soil, 391, 399–411.

Dechassa N, Merga M. 2022. Tea (Camellia sinensis) production, opportunities, challenges, and future prospects in Ethiopia. Advances in Agriculture, 2022, 1942666.

Deng W W, Fei Y, Wang S, Wan X C, Zhang Z Z, Hu X Y. 2013. Effect of shade treatment on theanine biosynthesis in Camellia sinensis seedlings. Plant Growth Regulation, 71, 295–299.

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.

Duan Y, Wang G, Liang L Y, Wang M H, Jiang J, Ma Y C, Zhu X J, Wu J, Fang W P. 2024a. Intercropping fruit trees in tea plantation improves soil properties and the formation of tea quality components. Plant Physiology and Biochemistry, 210, 108574.

Duan Y, Wang T, Lei X G, Cao Y, Liu L F, Zou Z W, Ma Y C, Zhu X J, Fang W P. 2024b. Leguminous green manure intercropping changes the soil microbial community and increases soil nutrients and key quality components of tea leaves. Horticulture Research, 11, uhae018.

Duan Y, Wang T, Zhang P X, Zhao X J, Jiang J, Ma Y C, Zhu X J, Fang W P. 2024c. The effect of intercropping leguminous green manure on theanine accumulation in the tea plant: A metagenomic analysis. Plant, Cell & Environment, 47, 1141–1159.

Fang C Y, Shen Y F, Wang F Y, Zhang J Y, Liu C, Luo F, Ye Y L. 2024. Combined transcriptomics and metabolomics to elucidate the underlying beneficial mechanisms of L-theanine in mitigating obesity-induced cardiac injury in rats. Journal of Functional Foods, 113, 106037.

Feng L, Yu Y C, Lin S J, Yang T Y, Chen Q, Liu L L, Sun J, Zheng P C, Zhang Z L, Wan X C. 2021. Tonoplast-localized theanine transporter CsCAT2 may mediate theanine storage in the root of tea plants (Camellia sinensis L.). Frontiers in Plant Science, 12, 797854.

Feng Y T, Sunderland T. 2023. Feasibility of tea/tree intercropping plantations on soil ecological service function in China. Agronomy, 13, 1548.

Fu X M, Cheng S H, Liao Y Y, Xu X L, Wang X C, Hao X Y, Xu P, Dong F, Yang Z Y. 2020. Characterization of L-theanine hydrolase in vitro and subcellular distribution of its specific product ethylamine in tea (Camellia sinensis). Journal of Agricultural and Food Chemistry, 68, 10842–10851.

Fung K F, Carr H P, Zhang J, Wong M H. 2008. Growth and nutrient uptake of tea under different aluminium concentrations. Journal of the Science of Food and Agriculture, 88, 1582–1591.

Godfray H C J, Poore J, Ritchie H. 2024. Opportunities to produce food from substantially less land. BMC Biology, 22, 138.

Hao T, Cui L J, Wang J Z, Lei Y R, Li W, Wang R M, Wang S K, Li J, Zhai X J, Zhang M Y, Zhao X S. 2025. Effects of low temperature on rhizosphere phosphate-mineralizing microbial populations in constructed wetlands. Journal of Environmental Management, 376, 124243.

Huang Z, Cui C H, Cao Y J, Dai J H, Cheng X Y, Hua S W, Wang W T, Duan Y, Petropoulos E, Wang H, Zhou L X, Fang W P, Zhong Z T. 2022. Tea plant–legume intercropping simultaneously improves soil fertility and tea quality by changing Bacillus species composition. Horticulture Research, 9, uhac046.

Hung Y, Chen P, Chen R L C, Cheng T. 2010. Sequential determination of tannin and total amino acid contents in tea for taste assessment by a fluorescent flow-injection analytical system. Food Chemistry, 118, 876–881.

Jia J Y, Zhang J Z, Li Y Z, Koziol L, Podzikowski L, Delgado-Baquerizo M, Wang G Z, Zhang J L. 2023. Relationships between soil biodiversity and multifunctionality in croplands depend on salinity and organic matter. Geoderma, 429, 116273.

Jiang Y H, Lin X Q, Wang H B, Xu Y N, Lin W X. 2025. Effect of intercropping with forage soybean on the rhizosphere soil nutrients of tea plants. Catena, 255, 109060.

Juneja L R, Chu D, 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 & Technology, 10, 199–204.

Kito M, Kokura H, Izaki J, Sasaoka K. 1968. Theanine, a precursor of the phloroglucinol nucleus of catechins in tea plants. Phytochemistry, 7, 599–603.

Lafleur B, Lalonde O, Labrecque M. 2017. First-rotation performance of five short-rotation willow cultivars on different soil types and along a large climate gradient. BioEnergy Research, 10, 158–166.

Lal R. 2009. Challenges and opportunities in soil organic matter research. European Journal of Soil Science, 60, 158–169.

Li N H, Gao D M, Zhou X G, Chen S C, Li C X, Wu F Z. 2020. Intercropping with potato-onion enhanced the soil microbial diversity of tomato. Microorganisms, 8, 834.

Li X F, Wang Z G, Bao X G, Sun J H, Yang S C, Wang P, Wang C B, Wu J P, Liu X R, Tian X L, Wang Yu, Li J P, Wang Y, Xia H Y, Mei P P, Wang X F, Zhao J H, Yu R P, Zhang W P, Che Z X, et al. 2021. Long-term increased grain yield and soil fertility from intercropping. Nature Sustainability, 4, 943–950.

Li X X, Zhang X H, Zhao Q S, Liao H. 2023. Genetic improvement of legume roots for adaption to acid soils. The Crop Journal, 11, 1022–1033.

Li Y H, Shi D Y, Li G H, Zhao B, Zhang J W, Liu P, Ren B Z, Dong S T. 2019. Maize/peanut intercropping increases photosynthetic characteristics, 13C-photosynthate distribution, and grain yield of summer maize. Journal of Integrative Agriculture, 18, 2219–2229.

Liu C G, Nie Y, Rao X, Tang J W, Siddique K H M. 2019. The effects of introducing Flemingia macrophylla to rubber plantations on soil water content and exchangeable cations. Catena, 172, 480–487.

Liu P F, Zhao Y K, Ma J N, Cao Y, Zhang M X, Yu J, Guan H B, Xing Y S, Wang X Q, Jia X. 2025. Impact of various intercropping modes on soil quality, microbial communities, yield and quality of Platycodon grandiflorum (Jacq.) A. DC. BMC Plant Biology, 25, 503.

Liu Y R, Lan X J, Hou H Q, Ji J H, Liu X M, Lv Z Z. 2024. Multifaceted ability of organic fertilizers to improve crop productivity and abiotic stress tolerance: Review and perspectives. Agronomy, 14, 1141.

Liu Z W, Wang Q, Wang T, Chen Q Y, He X F, Ma Y, Zhao M. 2025. Interplanting with leguminous plants improves theanine accumulation in tea shoots via the rhizosphere soil microecology. Industrial Crops and Products, 230, 121075.

Nowogrodzki A. 2019. How climate change might affect tea. Nature, 566, S10–S11.

Oldfield E E, Wood S A, Bradford M A. 2018. Direct effects of soil organic matter on productivity mirror those observed with organic amendments. Plant and Soil, 423, 363–373.

Osland M J, Gabler C A, Grace J B, Day R H, McCoy M L, McLeod J L, From A S, Enwright N M, Feher L C, Stagg C L, Hartley S B. 2018. Climate and plant controls on soil organic matter in coastal wetlands. Global Change Biology, 24, 5361–5379.

Ranjitkar S, Sujakhu N M, Lu Y, Wang Q, Wang M C, He J, Mortimer P E, Xu J C, Kindt R, Zomer R J. 2016. Climate modelling for agroforestry species selection in yunnan province, China. Environmental Modelling & Software, 75, 263–272.

Ruan J, Haerdter R, Gerendás J. 2010. Impact of nitrogen supply on carbon/nitrogen allocation: A case study on amino acids and catechins in green tea [Camellia sinensis (L.) O. Kuntze] plants. Plant Biology, 12, 724–734.

Ruan J Y, Ma L F, Yang Y J. 2012. Magnesium nutrition on accumulation and transport of amino acids in tea plants. Journal of the Science of Food and Agriculture, 92, 1375–1383.

Sahu N, Nayan R, Panda A, Varun A, Kesharwani R, Das P, Kumar A, Mallick S K, Mishra M M, Saini A, Aggarwal S P, Nayak S. 2025. Impact of changes in rainfall and temperature on production of Darjeeling tea in India. Atmosphere, 16, 1.

Shao S B, Li Z W, Ma X X, Cui J R, Zhu Y Q, Li Y P, Wu L K, Rensing C, Cai P M, Zhang J M, Li Q S. 2025. Enhancing tea plant growth and soil microbial ecology through intercropping tea plants with Ophiopogon japonicus. Plant and Soil, 513, 2807–2825.

Sun L T, Dong X, Wang Y, Maker G, Agarwal M, Ding Z T. 2022. Tea–soybean intercropping improves tea quality and nutrition uptake by inducing changes of rhizosphere bacterial communities. Microorganisms, 10, 2149.

Tilman D. 2020. Benefits of intensive agricultural intercropping. Nature Plants, 6, 604–605.

Tilman D, Balzer C, Hill J, Befort B L. 2011. Global food demand and the sustainable intensification of agriculture. Proceedings of the National Academy of Sciences of the United States of America, 108, 20260–20264.

Tilman D, Cassman K G, Matson P A, Naylor R, Polasky S. 2002. Agricultural sustainability and intensive production practices. Nature, 418, 671–677.

Vuong Q V, Bowyer M C, Roach P D. 2011. L-theanine: Properties, synthesis and isolation from tea. Journal of the Science of Food and Agriculture, 91, 1931–1939.

Wang D, Liu B J, Li F, Wang Z H, Hou J F, Cao R, Zheng Y Q, Yang W Q. 2025. Status and influential factors of soil nutrients and acidification in Chinese tea plantations: A meta-analysis. Soil, 11, 175–191.

Wang J F, Zhou Q, Lv Y L, Chen Z M. 2023. Effects of intercropping tea with landscape trees on ecosystem of tea garden and tea production. Acta Agriculturae Zhejiangensis, 35, 523–533. (in Chinese)

Wei H, Li H R, Wang Q, Xiang H M, Liu Z Q, Zhang J. 2025. Soil acidification alters C:N:P stoichiometry in the soil due to higher acid sensitivity of phosphorus. Environmental Science: Processes & Impacts, 27, 2094–2103.

Wen B, Zhang X L, Ren S, Duan Y, Zhang Y Y, Zhu X J, Wang Y H, Ma Y C, Fang W P. 2020. Characteristics of soil nutrients, heavy metals and tea quality in different intercropping patterns. Agroforestry Systems, 94, 963–974.

Wen X, Wu D L, Chen D J, Xu P R, Zhao T T, Chen S Y, Zhu Z H, Zhong H, Chen P. 2024. Soil organic matter and total nitrogen as key driving factors promoting the assessment of acid-base buffering characteristics in a tea (Camellia sinensis) plantation habitat. Environmental Monitoring and Assessment, 196, 596.

Wezel A, Casagrande M, Celette F, Vian J F, Ferrer A, Peigné J. 2014. Agroecological practices for sustainable agriculture. A review. Agronomy for Sustainable Development, 34, 1–20.

Wu H, Long X F, Geng Y F. 2023. Companion plants of tea: From ancient to terrace to forest. Plants, 12, 3061.

Wu Z H, Xue B, Wang S W, Xing X, Nuo M, Meng X, Wu M K, Jiang H, Ma H M, Yang M Y, Wei X S, Zhao G X, Tian P. 2024. Rice under dry cultivation–maize intercropping improves soil environment and increases total yield by regulating belowground root growth. Plants, 13, 2957.

Yan P, Wu L Q, Wang D H, Fu J Y, Shen C, Li X, Zhang L P, Zhang L, Fan L C, Han W Y. 2020. Soil acidification in Chinese tea plantations. Science of the Total Environment, 715, 136963.

Yan Z M, Zhong Y Z, Duan Y H, Chen Q H, Li F. 2020. Antioxidant mechanism of tea polyphenols and its impact on health benefits. Animal Nutrition, 6, 115–123.

Yang T Y, Xie Y X, Lu X, Yan X M, Wang Y, Ma J Z, Cheng X M, Lin S J, Bao S L, Wan X C, Lucas W J, Zhang Z L. 2021. Shading promoted theanine biosynthesis in the roots and allocation in the shoots of the tea plant (Camellia sinensis L.) cultivar shuchazao. Journal of Agricultural and Food Chemistry, 69, 4795–4803.

Yang X D, Ni K, Shi Y Z, Yi X Y, Zhang Q F, Fang L, Ma L F, Ruan J Y. 2018. Effects of long-term nitrogen application on soil acidification and solution chemistry of a tea plantation in China. Agriculture, Ecosystems & Environment, 252, 74–82.

Yang X D, Yi X Y, Ni K, Zhang Q F, Shi Y Z, Chen L B, Zhao Y Y, Zhang Y L, Ma Q X, Cai Y J, Ma L F, Ruan J Y. 2023. Patterns and abiotic drivers of soil organic carbon in perennial tea (Camellia sinensis L.) plantation system of China. Environmental Research, 237, 116925.

Ye J H, Wang H B, Yang X Y, Zhang Q, Li J Y, Jia X L, Kong X H, He H B. 2016. Autotoxicity of the soil of consecutively cultured tea plantations on tea (Camellia sinensis) seedlings. Acta Physiologiae Plantarum, 38, 195.

Ye J H, Wang Y H, Wang Y C, Hong L, Jia X L, Kang J Q, Lin S X, Wu Z Y, Wang H B. 2022. Improvement of soil acidification in tea plantations by long-term use of organic fertilizers and its effect on tea yield and quality. Frontiers in Plant Science, 13, 1055900.

Zhang J, Hu R B, Sreedasyam A, Garcia T M, Lipzen A, Wang M, Yerramsetty P, Liu D G, Ng V, Schmutz J, Cushman J C, Borland A M, Pasha A, Provart N J, Chen J G, Muchero W, Tuskan G A, Yang X H. 2020. Light-responsive expression atlas reveals the effects of light quality and intensity in Kalanchoë fedtschenkoi, a plant with crassulacean acid metabolism. GigaScience, 9, giaa018.

Zhang W J, Ni K, Long L Z, Ruan J Y. 2023. Nitrogen transport and assimilation in tea plant (Camellia sinensis): A review. Frontiers in Plant Science, 14, 1249202.

Zhao S X, Bai Y L, Jin Z, Long L, Diao W T, Chen W, Tan L Q, Tang Q, Tang D D. 2023. Effects of the combined application of nitrogen and selenium on tea quality and the expression of genes involved in nitrogen uptake and utilization in tea cultivar ‘Chuancha No.2.’ Agronomy, 13, 2997.

Zhao Y, Tian Y Q, Li X B, Song M H, Fang X X, Jiang Y, Xu X L. 2022. Nitrogen fixation and transfer between legumes and cereals under various cropping regimes. Rhizosphere, 22, 100546.

Zou Y, Shen F Y, Zhong Y N, Lv C N, Pokharel S S, Fang W P, Chen F J. 2022a. Impacts of intercropped maize ecological shading on tea foliar and functional components, insect pest diversity and soil microbes. Plants, 11, 1883.

Zou Y, Zhong Y N, Yu H, Pokharel S S, Fang W P, Chen F J. 2022b. Impacts of ecological shading by roadside trees on tea foliar nutritional and bioactive components, community diversity of insects and soil microbes in tea plantation. Biology, 11, 1800.