接种双色蜡蘑对马尾松根际土壤无机磷和活性铝含量的影响
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摘要
将双色蜡蘑(Laccaria bicolor)的3个株系(Lb 270、Lb S238A和Lb S238N)分别与马尾松种子拌匀,培养在酸性土壤中,以检测根际土壤无机磷和活性铝含量,并分析外生菌根对磷有效性和铝活性的作用。结果发现,与非菌根苗相比,接种双色蜡蘑显著促进马尾松幼苗生长和对磷、铝的吸收,而对根际土壤无机磷和活性铝含量的影响因菌株而异:Lb 270和Lb S238N显著降低pH、有效磷、全铝、交换态铝含量,提高羟基态铝含量,且Lb 270还显著降低铁磷和钙磷含量;Lb S238A显著提高pH、有效磷含量,减少铝磷、全铝、活性铝、交换态铝和腐殖酸铝含量。即接种双色蜡蘑提高了磷的生物有效性,且Lb 270和Lb S238N增加、Lb S238A减少铝的活性。因此,接种供试双色蜡蘑都能提高马尾松的抗铝性,主要在于其增加磷的生物有效性,而是否降低铝的活性取决于菌株特性。
【Objective】 In most soils, TP(total phosphorus) is generally plenty, but its availability to plants is very low. When the soil is getting acidified, soil Al is liable to get activated and then bound with HPO_4~(2-), H_2PO_4~-, and PO_4~(3-), turning them into hard-to-dissolve P, thus further lowering soil P availability. Some ectomycorrhizae(EM) are found to be able to deactivate soil Al, thus letting Al-complexes release bonded P in available form, for plants to take up. That is to say, some EMs could be introduced into acid soils to improve soil P availability and plant growth. In experiments of in vitro cultivation of EM fungi, some demonstrated their ability to raise P availability in soils neutral or slightly acidic, but little has been found in the literature validating such ability in the field. The present study aimed to assess effects of inoculation of EM fungi on inorganic P availability and Al activity in the rhizosphere soil of saplings planted in acid soil, and to explore mechanism of EM colonization improving host plants' resistance to Al toxicity. 【Method】 Germinating Pinus massoniana seeds, inoculated with three strains of Laccaria bicolor, namely Lb 270, Lb S238 A, and Lb S238 N, separately, were imbedded into pots of cold sandy yellow soil collected from a P. massoniana stand, and irrigated twice a day to maintain a proper moisture regime for 16 weeks under the natural conditions. At the end, comparison was made of Pinus massoniana saplings inoculated with EM fungi with those in the control(without inoculation). 【Result】 Results show that inoculation of L. bicolor significantly affected the contents of inorganic P and active Al in the rhizosphere soil of the saplings and the effects varied with the strain. Compared with the control, inoculation with any of the three L. bicolor strains significantly improved growth and P and Al, uptake of the saplings. In terms of the effect, the three strains exhibited an order of Lb S238 N > = Lb 270 > Lb S238 A. Inoculation with Lb 270 significantly decreased soil pH, the contents of available P, iron-bound P(Fe-P), calcium-bound P(Ca-P), total Al, and exchangeable Al, and the proportion of exchangeable Al in the fraction of labile Al, but increased the proportion of aluminum-bound P(Al-P) in the fraction of inorganic P, and the contents of labile Al or hydroxyl-Al and their proportions in total Al. Inoculation with Lb S238 A significantly reduced Al-P, the proportion of Al-P or Fe-P in the fraction of inorganic P, total Al, labile Al, exchangeable Al, organic-complexed Al, and the proportion of exchangeable Al and organic-complexed Al in the fraction of labile Al, but raised soil pH, the content of available P, the proportion of occluded-P(O-P) in the fraction of inorganic P, the proportion of labile Al in total Al, and proportions of hydroxyl-Al or acid-soluble Al in the fraction of labile Al. Inoculation with Lb S238 N significantly decreased soil pH, the contents of available P, total Al, exchangeable Al, and the proportions of exchangeable Al and organic-complexed Al in the fraction of labile Al, but increased the content of hydroxyl-Al, the proportion of labile Al in total Al and that of hydroxyl-Al in the fraction of active Al.【Conclusion】All the findings in this experiment demonstrate that inoculation with L. bicolor can improve the resistance of P. massoniana saplings to Al toxicity, which may be attributed to the ability of the fungi to enhance bio-availability of soil P, but their abilities to lower Al activity vary with strain of the fungi.
【Objective】 In most soils, TP(total phosphorus) is generally plenty, but its availability to plants is very low. When the soil is getting acidified, soil Al is liable to get activated and then bound with HPO_4~(2-), H_2PO_4~-, and PO_4~(3-), turning them into hard-to-dissolve P, thus further lowering soil P availability. Some ectomycorrhizae(EM) are found to be able to deactivate soil Al, thus letting Al-complexes release bonded P in available form, for plants to take up. That is to say, some EMs could be introduced into acid soils to improve soil P availability and plant growth. In experiments of in vitro cultivation of EM fungi, some demonstrated their ability to raise P availability in soils neutral or slightly acidic, but little has been found in the literature validating such ability in the field. The present study aimed to assess effects of inoculation of EM fungi on inorganic P availability and Al activity in the rhizosphere soil of saplings planted in acid soil, and to explore mechanism of EM colonization improving host plants' resistance to Al toxicity. 【Method】 Germinating Pinus massoniana seeds, inoculated with three strains of Laccaria bicolor, namely Lb 270, Lb S238 A, and Lb S238 N, separately, were imbedded into pots of cold sandy yellow soil collected from a P. massoniana stand, and irrigated twice a day to maintain a proper moisture regime for 16 weeks under the natural conditions. At the end, comparison was made of Pinus massoniana saplings inoculated with EM fungi with those in the control(without inoculation). 【Result】 Results show that inoculation of L. bicolor significantly affected the contents of inorganic P and active Al in the rhizosphere soil of the saplings and the effects varied with the strain. Compared with the control, inoculation with any of the three L. bicolor strains significantly improved growth and P and Al, uptake of the saplings. In terms of the effect, the three strains exhibited an order of Lb S238 N > = Lb 270 > Lb S238 A. Inoculation with Lb 270 significantly decreased soil pH, the contents of available P, iron-bound P(Fe-P), calcium-bound P(Ca-P), total Al, and exchangeable Al, and the proportion of exchangeable Al in the fraction of labile Al, but increased the proportion of aluminum-bound P(Al-P) in the fraction of inorganic P, and the contents of labile Al or hydroxyl-Al and their proportions in total Al. Inoculation with Lb S238 A significantly reduced Al-P, the proportion of Al-P or Fe-P in the fraction of inorganic P, total Al, labile Al, exchangeable Al, organic-complexed Al, and the proportion of exchangeable Al and organic-complexed Al in the fraction of labile Al, but raised soil pH, the content of available P, the proportion of occluded-P(O-P) in the fraction of inorganic P, the proportion of labile Al in total Al, and proportions of hydroxyl-Al or acid-soluble Al in the fraction of labile Al. Inoculation with Lb S238 N significantly decreased soil pH, the contents of available P, total Al, exchangeable Al, and the proportions of exchangeable Al and organic-complexed Al in the fraction of labile Al, but increased the content of hydroxyl-Al, the proportion of labile Al in total Al and that of hydroxyl-Al in the fraction of active Al.【Conclusion】All the findings in this experiment demonstrate that inoculation with L. bicolor can improve the resistance of P. massoniana saplings to Al toxicity, which may be attributed to the ability of the fungi to enhance bio-availability of soil P, but their abilities to lower Al activity vary with strain of the fungi.
引文
[1]Achat D L,Pousse N,Nico las M,et al.Soil properties controlling in organic phosphorus availability:General results from a national forest network and a global compilation of the literature.Biogeochemistry,2016,127(2):255-272
[2]Mimmo T,Ghizzi M,Cesco S,et al.Aluminumphosphate interactions in the rhizosphere of two bean species:Phaseolus lunatus L.and Phaseolus vulgaris L.Journal of the Science of Food and Agriculture,2013,93(15):3891-3896
[3]Opala P A,Okalebo J R,Othieno C O,et al.Effect of organic and inorganic phosphorus sources on maize yields in an acid soil in western Kenya.Nutrient Cycling in Agroecosystems,2010,86(3):317-329
[4]辜夕容,梁国仕,杨水平,等.接种双色蜡蘑对马尾松幼苗生长、养分和抗铝性的影响.林业科学,2005,41(4):199-203Gu X R,Liang G S,Yang S P,et al.Influences of Laccaria bicolor on the growth,nutrient uptake and aluminum resistance of Pinus massoniana seedlings(In Chinese).Scientia Silvae Sinicae,2005,41(4):199-203
[5]Tahara K,Norisada M,Tange T,et al.Ectomycorrhizal association enhances Al tolerance by inducing citrate secretion in Pinus densiflora.Soil Science and Plant Nutrition,2005,51(3):397-403
[6]Khosla B,Kaur H,Reddy M S.Influence of ectomycorrhizal colonization on the growth and mineral nutrition of Populus deltoides under aluminum toxicity.Journal of Plant Interaction,2009,4(2):93-99
[7]Ozasa T.A comprehensive analysis of the changes in organic acid metabolism of ectomycorrhizal fungus Laccaria bicolor in response to aluminum salts.Sustainable Humanosphere:Bulletin of Research Institute for Sustainable Humanosphere Kyoto University,2013:33
[8]Zhang L,Wang M X,Li H,et al.Mobilization of inorganic phosphorus from soils by ectomycorrhizal fungi.Pedosphere,2014,24(5):683-689
[9]Adeleke R,Nwangburuka C,Oboirien B.Origins,roles and fate of organic acids in soils:A review.South African Journal of Botany,2017,108:393-406
[10]王明霞,袁玲,周志峰,等.铝对外生菌根真菌草酸分泌及氮磷钾吸收的影响.林业科学,2012,48(2):82-88Wang M X,Yuan L,Zhou Z F,et al.Efflux of oxalate and uptake of nitrogen,phosphorus and potassium by ectomycorrhizal fungi isolates in vitro in response to aluminum stress(In Chinese).Scientia Silvae Sinicae,2012,48(2):82-88
[11]李华,黄建国,袁玲.铝和锰对外生菌根真菌生长、养分吸收及分泌作用的影响.环境科学,2013,34(1):315-320Li H,Huang J G,Yuan L.Influence of aluminum and manganese on growth,nutrient uptake and the efflux by ectomycorrhizal fungi(In Chinese).Chinese Journal of Environmental Science,2013,34(1):315-320
[12]辜夕容,黄建国.铝对外生菌根真菌草酸分泌及磷、钾、铝吸收的影响.生态学报,2010,30(2):357-363Gu X R,Huang J G.Effect of aluminum on growth,oxalate exudation,and uptake of aluminum,phosphorus and potassium by ectomycorrhizal fungi in vitro(In Chinese).Acta Ecologica Sinica,2010,30(2):357-363
[13]吴雪莲,周倩,辜夕容,等.西南林区3株外生菌根真菌对酸铝胁迫的营养响应与抗铝性.西南大学学报(自然科学版),2015,37(8):13-19Wu X L,Zhou Q,Gu X R,et al.Resistance and nutritional response to aluminum toxicity of three ectomycorrhizal fungi from forest soils in Southwest China(In Chinese).Journal of Southwest University(Natural Science Edition),2015,37(8):13-19
[14]Heim A,Brunner I,Frossard E,et al.Aluminum effects on Picea abies at low solution concentrations.Soil Science Society of America Journal,2003,67(3):895-898
[15]吕焕哲,王凯荣,谢小立.土地利用方式与坡位土壤活性铝形态特征分析.水土保持学报,2007,21(1):172-175LüH Z,Wang K R,Xie X L.Character of soil aluminum forms under different land use and slope position(In Chinese).Journal of Soil and Water Conservation,2007,21(1):172-175
[16]Sholl L V,Kuyper T W,Smits M M,et al.Rockeating mycorrhizas:Their role in plant nutrition and biogeochemical cycles.Plant and Soil,2008,303(1):35-47
[17]周垂帆,林静雯,李莹,等.草甘膦对土壤磷形态及有效性的影响.西北林学院学报,2016,31(6):71-77Zhou C F,Lin J W,Li Y,et al.Effects of glyphosate on inorganic phosphorus transformation in soil(In Chinese).Journal of Northwest Forestry University,2016,31(6):71-77
[18]国家林业局.森林植物与森林枯枝落叶层全硅、铁、铝、钙、镁、钾、钠、磷、硫、锰、铜、锌的测定(LY/T 1270-1999).北京:中国标准出版社,1999:279-294State Forestry Administration of China.Determination of total silica,iron,aluminum,calcium,magnesium,potassium,sodium,phosphorus,sulphur,manganese,copper and zinc in forest plant and forest floor.Beijing:China Standard Press,1999:279-294
[19]鲍士旦.土壤农化分析.第3版.北京:中国农业出版社,2000:74-165Bao S D.Soil and agricultural chemistry analysis(In Chinese).3rd ed.Beijing:China Agriculture Press,2000:74-165
[20]黄衍初,曲长菱.土壤中铝的溶出及形态研究.环境科学,1996,17(1):57-59Huang Y C,Qu C L.Studies on the leaching and species of aluminum in soil(In Chinese).Chinese Journal of Environmental Science,1996,17(1):57-59
[21]Huang J G,Lapeyrie F.Ability of ectomycorrhizal fungus Laccaria bicolor S238N to increase the growth of Douglas fir seedlings and their phosphorus and potassium uptake.Pedosphere,1996,6(3):217-224
[22]刘进法,夏仁学,王明元,等.丛枝菌根促进植物根系吸收难溶态磷的研究进展(综述).亚热带植物科学,2007,36(4):62-66Liu J F,Xia R X,Wang M Y,et al.Recent advances in phosphorus acquisition from non-labile sources in plants with arbuscular mycorrhizal interaction(In Chinese).Subtropical Plant Science,2007,36(4):62-66
[23]Harrold S A,Tabatabai M A.Release of inorganic phosphorus from soils by low-molecular-weight organic acids.Communications in Soil Science and Plant Analysis,2006,37(9):1233-1245
[24]龚松贵,王兴祥,张桃林,等.低分子量有机酸对红壤无机磷活化的作用.土壤学报,2010,47(4):692-697Gong S G,Wang X X,Zhang T L,et al.Release of inorganic phosphorus from red soils induced by low molecular weight organic acids(In Chinese).Acta Pedologica Sinica,2010,47(4):692-697
[25]王玉书,刘海,袁玲.空心莲子草根系分泌物对无机磷细菌的负化感效应.土壤学报,2017,54(6):1486-1496Wang Y S,Liu H,Yuan L.Negative allelopathic effects of root exudate of Alternanthera philoxeroides on growth and phosphate dissolution of inorganic phosphorus bacteria(In Chinese).Acta Pedologica Sinica,2017,54(6):1486-1496
[26]Ray P,Adholeya A.Correlation between organic acid exudation and metal uptake by ectomycorrhizal fungi grown on pond ash in vitro.Biometals,2009,22(2):275-281
[27]Ahonen-Jonnarth U,Hees P A W V,Lundstrom US,et al.Organic acids produced by mycorrhizal Pinus sylvestris exposed to elevated aluminum and heavy metal concentrations.New Phytologist,2000,146(3):557-567
[28]王斌,张慧玲,朱晓芳,等.水稻品种间铝耐性差异的探究.土壤学报,2017,54(4):958-966Wang B,Zhang H L,Zhu X F,et al.Differences in aluminum tolerance between rice varieties(In Chinese).Acta Pedologica Sinica,2017,54(4):958-966
[29]刘鹏,金婷婷,徐根娣,等.大豆根系分泌物对根际土壤铝形态的影响.浙江师范大学学报(自然科学版),2008,31(4):445-451Liu P,Jin T T,Xu G D,et al.The effect of soybean’s root exudate on the forms of aluminum of rhizosphere soil(In Chinese).Journal of Zhejiang Normal University(Natural Sciences),2008,31(4):445-451
[30]张俊思,袁大刚,付宏阳,等.EGCG溶液浓度与酸碱度对黄壤Al、Fe和Mn形态的影响.土壤学报,2017,54(4):905-916Zhang J S,Yuan D G,Fu H Y,et al.Effect of EGCG on Al,Fe and Mn in yellow soil relative to concentration and pH(In Chinese).Acta Pedologica Sinica,2017,54(4):905-916
[2]Mimmo T,Ghizzi M,Cesco S,et al.Aluminumphosphate interactions in the rhizosphere of two bean species:Phaseolus lunatus L.and Phaseolus vulgaris L.Journal of the Science of Food and Agriculture,2013,93(15):3891-3896
[3]Opala P A,Okalebo J R,Othieno C O,et al.Effect of organic and inorganic phosphorus sources on maize yields in an acid soil in western Kenya.Nutrient Cycling in Agroecosystems,2010,86(3):317-329
[4]辜夕容,梁国仕,杨水平,等.接种双色蜡蘑对马尾松幼苗生长、养分和抗铝性的影响.林业科学,2005,41(4):199-203Gu X R,Liang G S,Yang S P,et al.Influences of Laccaria bicolor on the growth,nutrient uptake and aluminum resistance of Pinus massoniana seedlings(In Chinese).Scientia Silvae Sinicae,2005,41(4):199-203
[5]Tahara K,Norisada M,Tange T,et al.Ectomycorrhizal association enhances Al tolerance by inducing citrate secretion in Pinus densiflora.Soil Science and Plant Nutrition,2005,51(3):397-403
[6]Khosla B,Kaur H,Reddy M S.Influence of ectomycorrhizal colonization on the growth and mineral nutrition of Populus deltoides under aluminum toxicity.Journal of Plant Interaction,2009,4(2):93-99
[7]Ozasa T.A comprehensive analysis of the changes in organic acid metabolism of ectomycorrhizal fungus Laccaria bicolor in response to aluminum salts.Sustainable Humanosphere:Bulletin of Research Institute for Sustainable Humanosphere Kyoto University,2013:33
[8]Zhang L,Wang M X,Li H,et al.Mobilization of inorganic phosphorus from soils by ectomycorrhizal fungi.Pedosphere,2014,24(5):683-689
[9]Adeleke R,Nwangburuka C,Oboirien B.Origins,roles and fate of organic acids in soils:A review.South African Journal of Botany,2017,108:393-406
[10]王明霞,袁玲,周志峰,等.铝对外生菌根真菌草酸分泌及氮磷钾吸收的影响.林业科学,2012,48(2):82-88Wang M X,Yuan L,Zhou Z F,et al.Efflux of oxalate and uptake of nitrogen,phosphorus and potassium by ectomycorrhizal fungi isolates in vitro in response to aluminum stress(In Chinese).Scientia Silvae Sinicae,2012,48(2):82-88
[11]李华,黄建国,袁玲.铝和锰对外生菌根真菌生长、养分吸收及分泌作用的影响.环境科学,2013,34(1):315-320Li H,Huang J G,Yuan L.Influence of aluminum and manganese on growth,nutrient uptake and the efflux by ectomycorrhizal fungi(In Chinese).Chinese Journal of Environmental Science,2013,34(1):315-320
[12]辜夕容,黄建国.铝对外生菌根真菌草酸分泌及磷、钾、铝吸收的影响.生态学报,2010,30(2):357-363Gu X R,Huang J G.Effect of aluminum on growth,oxalate exudation,and uptake of aluminum,phosphorus and potassium by ectomycorrhizal fungi in vitro(In Chinese).Acta Ecologica Sinica,2010,30(2):357-363
[13]吴雪莲,周倩,辜夕容,等.西南林区3株外生菌根真菌对酸铝胁迫的营养响应与抗铝性.西南大学学报(自然科学版),2015,37(8):13-19Wu X L,Zhou Q,Gu X R,et al.Resistance and nutritional response to aluminum toxicity of three ectomycorrhizal fungi from forest soils in Southwest China(In Chinese).Journal of Southwest University(Natural Science Edition),2015,37(8):13-19
[14]Heim A,Brunner I,Frossard E,et al.Aluminum effects on Picea abies at low solution concentrations.Soil Science Society of America Journal,2003,67(3):895-898
[15]吕焕哲,王凯荣,谢小立.土地利用方式与坡位土壤活性铝形态特征分析.水土保持学报,2007,21(1):172-175LüH Z,Wang K R,Xie X L.Character of soil aluminum forms under different land use and slope position(In Chinese).Journal of Soil and Water Conservation,2007,21(1):172-175
[16]Sholl L V,Kuyper T W,Smits M M,et al.Rockeating mycorrhizas:Their role in plant nutrition and biogeochemical cycles.Plant and Soil,2008,303(1):35-47
[17]周垂帆,林静雯,李莹,等.草甘膦对土壤磷形态及有效性的影响.西北林学院学报,2016,31(6):71-77Zhou C F,Lin J W,Li Y,et al.Effects of glyphosate on inorganic phosphorus transformation in soil(In Chinese).Journal of Northwest Forestry University,2016,31(6):71-77
[18]国家林业局.森林植物与森林枯枝落叶层全硅、铁、铝、钙、镁、钾、钠、磷、硫、锰、铜、锌的测定(LY/T 1270-1999).北京:中国标准出版社,1999:279-294State Forestry Administration of China.Determination of total silica,iron,aluminum,calcium,magnesium,potassium,sodium,phosphorus,sulphur,manganese,copper and zinc in forest plant and forest floor.Beijing:China Standard Press,1999:279-294
[19]鲍士旦.土壤农化分析.第3版.北京:中国农业出版社,2000:74-165Bao S D.Soil and agricultural chemistry analysis(In Chinese).3rd ed.Beijing:China Agriculture Press,2000:74-165
[20]黄衍初,曲长菱.土壤中铝的溶出及形态研究.环境科学,1996,17(1):57-59Huang Y C,Qu C L.Studies on the leaching and species of aluminum in soil(In Chinese).Chinese Journal of Environmental Science,1996,17(1):57-59
[21]Huang J G,Lapeyrie F.Ability of ectomycorrhizal fungus Laccaria bicolor S238N to increase the growth of Douglas fir seedlings and their phosphorus and potassium uptake.Pedosphere,1996,6(3):217-224
[22]刘进法,夏仁学,王明元,等.丛枝菌根促进植物根系吸收难溶态磷的研究进展(综述).亚热带植物科学,2007,36(4):62-66Liu J F,Xia R X,Wang M Y,et al.Recent advances in phosphorus acquisition from non-labile sources in plants with arbuscular mycorrhizal interaction(In Chinese).Subtropical Plant Science,2007,36(4):62-66
[23]Harrold S A,Tabatabai M A.Release of inorganic phosphorus from soils by low-molecular-weight organic acids.Communications in Soil Science and Plant Analysis,2006,37(9):1233-1245
[24]龚松贵,王兴祥,张桃林,等.低分子量有机酸对红壤无机磷活化的作用.土壤学报,2010,47(4):692-697Gong S G,Wang X X,Zhang T L,et al.Release of inorganic phosphorus from red soils induced by low molecular weight organic acids(In Chinese).Acta Pedologica Sinica,2010,47(4):692-697
[25]王玉书,刘海,袁玲.空心莲子草根系分泌物对无机磷细菌的负化感效应.土壤学报,2017,54(6):1486-1496Wang Y S,Liu H,Yuan L.Negative allelopathic effects of root exudate of Alternanthera philoxeroides on growth and phosphate dissolution of inorganic phosphorus bacteria(In Chinese).Acta Pedologica Sinica,2017,54(6):1486-1496
[26]Ray P,Adholeya A.Correlation between organic acid exudation and metal uptake by ectomycorrhizal fungi grown on pond ash in vitro.Biometals,2009,22(2):275-281
[27]Ahonen-Jonnarth U,Hees P A W V,Lundstrom US,et al.Organic acids produced by mycorrhizal Pinus sylvestris exposed to elevated aluminum and heavy metal concentrations.New Phytologist,2000,146(3):557-567
[28]王斌,张慧玲,朱晓芳,等.水稻品种间铝耐性差异的探究.土壤学报,2017,54(4):958-966Wang B,Zhang H L,Zhu X F,et al.Differences in aluminum tolerance between rice varieties(In Chinese).Acta Pedologica Sinica,2017,54(4):958-966
[29]刘鹏,金婷婷,徐根娣,等.大豆根系分泌物对根际土壤铝形态的影响.浙江师范大学学报(自然科学版),2008,31(4):445-451Liu P,Jin T T,Xu G D,et al.The effect of soybean’s root exudate on the forms of aluminum of rhizosphere soil(In Chinese).Journal of Zhejiang Normal University(Natural Sciences),2008,31(4):445-451
[30]张俊思,袁大刚,付宏阳,等.EGCG溶液浓度与酸碱度对黄壤Al、Fe和Mn形态的影响.土壤学报,2017,54(4):905-916Zhang J S,Yuan D G,Fu H Y,et al.Effect of EGCG on Al,Fe and Mn in yellow soil relative to concentration and pH(In Chinese).Acta Pedologica Sinica,2017,54(4):905-916