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Figure 1.
Fluoride absorption by tea plants. Tea plants absorb fluoride from the atmosphere, soil, and water. Fluoride in the atmosphere is absorbed through the stomata or cuticle of the leaf epidermis. Fluoride ions and fluoride–aluminum complexes in soil and water are absorbed by the roots.
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Figure 2.
Fluoride transportation in tea plants. Fluoride absorbed by the leaves is transferred to the leaf tips and edges, chelates with metal ions, and the complexes are deposited in the leaves. Fluoride can be transferred from old leaves to new tips. Fluoride from soil and water can form complexes with organic acids and aluminum, which are stored in the leaves of tea plants. Fluoride from soil and water can also be individually absorbed and transported for storage in the leaves.
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Figure 3.
Mechanisms of fluoride tolerance in tea plants. (A) Metabolites in tea plants reduce the toxic effect of fluoride. Cell wall macromolecular components such as pectin, lignin, cellulose, hemicellulose, polysaccharide, and proteins chelate fluoride. The contents of some metabolites increase during adaptation of tea plants to fluoride. (B) Complexation of cations (Al3+, Ca2+, and Mg2+) with fluoride in tea plants. (C) Roles of the antioxidant system of tea plant in fluoride tolerance. Increases in the activity/abundance of antioxidant enzymes and in the ASA-GSH cycle reduce intracellular levels of reactive oxygen species, leading to increased fluoride tolerance. (D) Roles of transporters in fluoride tolerance. Transporter proteins transport fluoride into the vacuole, and this compartmentalization reduces damage to enzymes and organelles. The CsFEX, CsCLC, and CsABC transporters efflux fluoride from cells, thereby reducing its toxic effects. POD: Peroxidase, CAT: Catalase, SOD: Superoxide dismutase, APX: Ascorbate peroxidase, GR: Glutathione reductase, DHAR: Dehydroascorbate reductase, ABA: Abscisic acid, GA: Gibberellic acid, GHs: Glycoside hydrolases, ASA-GSH: Antioxidant system and the ascorbate-glutathione, ROS: Reactive oxygen species.
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Figure 4.
Effects of fluoride stress on tea plant growth and tea quality. Fluoride at low concentrations increases the chlorophyll content, photosynthetic rate, and quality-related metabolites in tea plants; and increases the activity of the antioxidant system and the ASA-GSH cycle to remove reactive oxygen species (ROS). Fluoride at high concentrations that exceed the tolerance limit of tea plants decreases the scavenging capacity of the antioxidant system and the ASA-GSH cycle, resulting in ROS accumulation. In addition, damage to chloroplast thylakoid membranes and decreases in chlorophyll content lead to decreases in the photosynthetic rate, stomatal conductance, and carbon assimilation capacity, resulting in decreased biomass and decreased content of quality-related metabolites, as well as leaf yellowing, leaf abscission, and even plant death. POD: Peroxidase, CAT: Catalase, SOD: Superoxide dismutase, APX: Ascorbate peroxidase, GR: Glutathione reductase, DHAR: Dehydroascorbate reductase, ASA-GSH: Antioxidant system and the ascorbate-glutathione, ROS: Reactive oxygen species.
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Figure 5.
Defluoridation measures for tea plants. (A) Breeding low-fluoride varieties of tea plants. (B) Improving management measures during tea plant cultivation. (C) Improving tea processing technologies. (D) Appropriate brewing methods to prepare tea infusions.
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Cultivars Province Parts Treatment Years Content (mg/kg) Ref. Liannandaye Sichuan Old leaves Drying at 80 °C and boiling water extraction 2006−2007 1,150.79 ± 4.86 [107] Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 1,296.66 ± 12.84 [110] Yuenandaye Old leaves Drying at 80 °C and boiling water extraction 2006−2007 1,352.89 ± 12.69 [107] Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 1,560.36 ± 27.10 [110] Chenxi NO.4 Old leaves Drying at 80 °C and boiling water extraction 2006−2007 1,865.61 ± 7.46 [107] Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 1,954.93 ± 10.96 [110] Meizhan Old leaves Drying at 80 °C and boiling water extraction 2006−2007 2,180.13 ± 14.42 [107] Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 1,732.2 ± 41.2 [110] Zhejiang Mature leaves Drying at 80 °C and nitric acid extraction 2002 2,015.48 ± 29.99 [106] Fudingdabai Sichuan Old leaves Drying at 80 °C and boiling water extraction 2006−2007 249.64 ± 24.3 [107] Guizhou One bud and
five leavesDrying at 80 °C and hydrochloric acid extraction 2010 1,612.3 ± 43.1 [109] Fudingdabai Zhejiang Mature leaves Drying at 80 °C and nitric acid extraction 2002 137.1 ± 2.1 [106] Fujian Old leaves Drying at 80 °C and hydrochloric acid extraction 2010 282.1 [111] Hunan One bud and
five leavesSteaming and boiling water extraction 2011 2,232.05 ± 85.52 [51] Zhuyeqi Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,750.16 ± 11.37 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 125.4 [110] Hunan One bud and
five leavesSteaming and boiling water extraction 2011 2,330.74 ± 31.39 [51] Fujianshuixian Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,548.18 ± 40.97 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 103.7 ± 1.5 [110] Fujian Old leaves Drying at 80 °C and hydrochloric acid extraction 2010 1,150.79 ± 4.86 [111] Huangyeshuixian Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,424.70 ± 18.85 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 2,950.80 ± 27.73 [110] Qianmei 701 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,522.01 ± 45.33 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,693.09 ± 35.12 [110] Guizhou One bud and
five leavesDrying at 80 °C and hydrochloric acid extraction 2010 389.95 ± 32.18 [109] Guizhou Old leaves Dry samples and hydrochloric acid extraction 2011 2,142.26 ± 16.30 [113] Mingshan 130 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,564.78 ± 51.22 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,036.13 ± 31.25 [110] Mengshan 9 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,647.31 ± 70.89 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,436.55 ± 20.21 [110] Yinghong NO. 2 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,669.02 ± 799.95 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,364.53 ± 51.72 [110] Mengshan 11 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,695.21 ± 59.89 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,582.83 ± 9.73 [110] Mingshan 311 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,716.22 ± 42.21 [107] Donghuzao 2,731.20 ± 20.78 Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,107.27 ± 54.91 [110] Hainandaye Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,746.82 ± 39.71 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 2,961.53 ± 29.94 [110] Qianmei 502 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,878.23 ± 76.94 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,881.51 ± 16.48 [110] Guizhou One bud and
five leavesDrying at 80 °C and hydrochloric acid extraction 2010 389.95 ± 30.2 [109] Old leaves Dry samples and hydrochloric acid extraction 2011 3,260.48 ± 32.12 [113] Zisun Ya'an and
surroundingsDrying at 80 °C and boiling water extraction 2006−2007 2,904.13 ± 35.40 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,140.80 ± 42.86 [110] Zhejiang Drying at 80 °C and nitric acid extraction 2002 1,742.7 ± 43.2 [106] Qianmei 303 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,918.13 ± 46.79 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 4,029.11 ± 81.86 [110] Guizhou One bud and
five leavesDrying at 80 °C and hydrochloric acid extraction 2010 199.74 ± 16.6 [109] Old leaves Dry samples and hydrochloric acid extraction 2011 2,972.79 ± 169.82 [113] Anxishuixian Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 2,924.33 ± 41.39 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,454.68 ± 26.29 [110] Longjing 43 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 3,152.73 ± 27.70 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 4,437.79 ± 26.14 [110] Zhejiang Drying at 80 °C and nitric acid extraction 2002 1,377.1 ± 37.0 [106] Fujian Old leaves Drying at 80 °C and hydrochloric acid extraction 2010 116.2 ± 0.9 [111] Shuyong 307 Ya'an and
surroundingsDrying at 80 °C and boiling water extraction 2006−2007 3,223.55 ± 151.43 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 4,296.52 ± 54.98 [110] Zhenghedabaicha Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 3,295.74 ± 27.55 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 3,876.58 ± 21.09 [110] Zhejiang Drying at 80 °C and nitric acid extraction 2002 1,373.0 ± 41.9 [106] Taiwandaye Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 3,363.59 ± 456.39 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 4,739.89 ± 58.59 [110] Qianmei 419 Ya'an and
surroundingsOld leaves Drying at 80 °C and boiling water extraction 2006−2007 3,518.15 ± 76.19 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 4,541.43 ± 28.91 [110] Guizhou One bud and
five leavesDrying at 80 °C and hydrochloric acid extraction 2010 133.70 ± 11.2 [109] Old leaves Dry samples and hydrochloric acid extraction 2011 2,370.47 ± 11.43 [113] Mengshan23 Ya'an and
surroundingsDrying at 80 °C and boiling water extraction 2006−2007 3,625.11 ± 86.07 [107] Ya'an Mature leaves Drying at 70 °C and hydrochloric acid extraction 2006 4,469.25 ± 40.85 [110] Sichuan group species 2,782.59 ± 146.46 Meitantaicha Guizhou One bud and
five leavesDrying at 80 °C and hydrochloric acid extraction 2010 272.93 ± 27.3 [109] Qianmei 101 Drying at 80 °C and hydrochloric acid extraction 2010 175.07 ± 13.5 Dry samples and hydrochloric acid extraction 2011 3,218.33 ± 57.91 [113] Qianmei 601 Drying at 80 °C and hydrochloric acid extraction 2010 267.11 ± 26.31 [109] Dry samples and hydrochloric acid extraction 2011 2,823.02 ± 73.36 [113] Qianmei 809 Drying at 80 °C and hydrochloric acid extraction 2010 521.48 ± 50.32 [109] Dry samples and hydrochloric acid extraction 2011 2,327.91 ± 83.17 [113] Qianmei 308 Drying at 80 °C and hydrochloric acid extraction 2010 326.88 ± 29.3 [109] Dry samples and hydrochloric acid extraction 2011 3,432.86 ± 159.4 [113] Qianmei 415 Drying at 80 °C and hydrochloric acid extraction 2010 186.95 ± 14.2 [109] Dry samples and hydrochloric acid extraction 2011 5,090.83 ± 69.56 [113] Qiancha NO. 7 Drying at 80 °C and hydrochloric acid extraction 2010 218.81 ± 18.7 [109] Qianfu NO. 4 136.82 ± 11.6 Dry samples and hydrochloric acid extraction 2011 3,066.49 ± 86.35 [113] Guiyucha NO. 8 Drying at 80 °C and hydrochloric acid extraction 2010 191.03 ± 18.6 [109] Dry samples and hydrochloric acid extraction 2011 2,882.94 ± 195.73 [113] Pingyangtezao Drying at 80 °C and hydrochloric acid extraction 2010 125.02 ± 12.1 [109] Yuanxiaolv 244.32 ± 20.5 Nongkangzao 133.70 ± 12.3 Mingshan 213 106.98 ± 6.74 Mingke NO. 4 195.29 ± 16.8 Maolv 174.73 ± 15.8 Qianmei 412 Old leaves Dry samples and hydrochloric acid extraction 2011 3,396.92 ± 31.61 [113] Meitantaicha 2011 3,085.83 ± 101.9 Zhenong 138 Zhejiang Mature leaves Drying at 80 °C and nitric acid extraction 2002 805.7 ± 6.0 [106] Zhenong 12 1,041.2 ± 23.3 Shuigu 1,123.2 ± 33.5 Hanlv 1,152.4 ± 2.4 Zhuzhichun 1,248.2 ± 2.3 Lvyafoshou 1,298.1 ± 2.0 Zhenong 139 1,322.4 ± 40.7 Shuixian 1,323.5 ± 36.1 Biyun 1,400.9 ± 0.6 Soubei 1,487.6 ± 29.7 Maoxie 1,487.7 ± 31.0 Anhui NO. 9 1,489.7 ± 40.0 Zhenong 113 1,492.7 ± 43.5 Yingshuang 1,509.9 ± 7.2 Zhenong 25 1,521.2 ± 3.2 Ribenzhong 1,543.8 ± 33.9 Yunqi 1,549.3 ± 46.9 Zhenong 23 1,576.7 ± 11.3 Huangyezao 1,606.7 ± 40.5 Jinshi 1,662.4 ± 42.4 Pingyun 1,676.6 ± 44.6 Zhenong 21 1,678.8 ± 49.6 Jinfeng 1,705.2 ± 10.3 Jiukeng 1,779.2 ± 5.0 Juhuachun 1,993.4 ± 14.5 Wuniuzao 2,163.2 ± 15.8 Fujian Old leaves Drying at 80 °C and hydrochloric acid extraction 2010 98.0 ± 1.3 [111] Jinguanyin All leaves Drying at 80 °C and nitric acid extraction 2014−2015 536.49 ± 10.41 [112] Dangui 2,598.87 ± 24.12 Old leaves Drying at 80 °C and hydrochloric acid extraction 2010 145.3 ± 0.2 [111] Jinmudan All leaves Drying at 80 °C and nitric acid extraction 2014−2015 1,030.21 ± 36.52 [112] Ruixiang Drying at 80 °C and nitric acid extraction 1,315.64 ± 21.56 Xiapu yuanxiao Old leaves Drying at 80 °C and hydrochloric acid extraction 2010 124.6 ± 3.0 [111] Jinxuan 103.7 ± 3.5 Fuandabai 103.0 ± 1.0 Fuyun NO. 7 104.0 ± 1.1 Fuyun NO. 6 131.6 ± 1.8 Fudingdahao 118.0 ± 2.4 Zaochunhao 107.7 ± 2.8 Xiapu chunbolv 99.2 ± 1.2 BaijiguanF1 102.1 ± 1.1 Huanguanyin Old leaves Drying at 80 °C and hydrochloric acid extraction 99.2 ± 1.3 Foxiang NO. 1 Yunnan One bud and
four leavesDrying at 60 °C and hydrochloric acid extraction 2011 155.00 ± 6.94 [108] Foxiang NO. 2 214.30 ± 5. 94 Foxiang NO. 4 219. 50 ± 7. 32 Foxiang NO. 5 190.70 ± 4.09 Yunkang NO. 10 121. 30 ± 5. 81 Yunkang NO. 14 198.50 ± 8.49 Yuncha NO. 1 135.10 ± 4.74 Baihaozao Hunan One bud and
five leavesSteaming and boiling water extraction 2011 113.2 [51] Bixiangzao 121.4 Taoyuandaye 177.7 Yulv 165.9 Jianbohuang NO.13 168.5 Gaoyaqi 162.8 Table 1.
Fluoride content difference of different tea cultivars.
Figures
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Tables
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