安吉白茶阶段性白化过程中光合特性的变化
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摘要
为阐明安吉白茶阶段性白化过程中光合特性的变化,本研究以特殊茶树资源安吉白茶为材料,研究其阶段性白化过程中叶绿体超微结构、光合作用、光合色素和生化成分的变化。结果表明,安吉白茶白化阶段叶绿体发育受阻,已发育的叶绿体超微结构被破坏,复绿阶段则逐渐恢复正常;净光合速率、气孔导度和蒸腾作用在阶段性白化过程中整体呈上升趋势,而在白化期显著下降;胞间CO2浓度在阶段性白化过程中整体呈下降趋势,而在白化期有所升高;光合色素和茶多酚含量在阶段性白化过程中整体逐渐上升,而在白化期显著降低;总氨基酸和茶氨酸含量在阶段性白化过程中整体逐渐下降,而在白化期显著升高。为安吉白茶叶色突变和高氨基酸含量形成机制研究提供了一定的理论基础。
In order to elucidate the changes of photosynthetic characteristics during the periodic albinism of Anji white tea, this study took Anji white tea(special tea tree resources) as materials, and investigated the changes of chloroplast ultrastructure, photosynthesis, photosynthetic pigments and biochemical components in Anji white tea during periodic albinism. The results showed that the development of chloroplast was hindered and the ultrastructure of chloroplast was destroyed in the albinistic stage, but it gradually returned to normal in the re-green stage. The net photosynthetic rate, stomatal conductance, and transpiration gradually increased during periodic albinism, while decreased significantly in the albinistic stage. The concentration of intercellular CO2 showed a downward trend during periodic albinism, while slightly increased in the albinistic stage. The photosynthetic pigments and tea polyphenols were significantly increased during periodic albinism, while significantly decreased in the albinistic stage. The total amino acid and theanine content gradually decreased during the periodic albinism,while significantly increased in the albinistic stage. It might provide a theoretical basis for the research on the for mation mechanism of color mutation and high amino acid content in Anji white tea.
In order to elucidate the changes of photosynthetic characteristics during the periodic albinism of Anji white tea, this study took Anji white tea(special tea tree resources) as materials, and investigated the changes of chloroplast ultrastructure, photosynthesis, photosynthetic pigments and biochemical components in Anji white tea during periodic albinism. The results showed that the development of chloroplast was hindered and the ultrastructure of chloroplast was destroyed in the albinistic stage, but it gradually returned to normal in the re-green stage. The net photosynthetic rate, stomatal conductance, and transpiration gradually increased during periodic albinism, while decreased significantly in the albinistic stage. The concentration of intercellular CO2 showed a downward trend during periodic albinism, while slightly increased in the albinistic stage. The photosynthetic pigments and tea polyphenols were significantly increased during periodic albinism, while significantly decreased in the albinistic stage. The total amino acid and theanine content gradually decreased during the periodic albinism,while significantly increased in the albinistic stage. It might provide a theoretical basis for the research on the for mation mechanism of color mutation and high amino acid content in Anji white tea.
引文
Abe T.,Matsuyama T.,Sekido S.,Yamaguchi I.,Yoshida S.,and Kameya T.,2002,Chlorophyll-deficient mutants of rice demonstrated the deletion of a DNA fragment by heavy-ion irradiation,Journal of Radiation Research,43(4):S157-S161
Asakura Y.,Hi rohashi T.,Kikuchi S.,Belcher S.,Osborne E.,Yano S.,Terashima I.,Barkan A.,and Nakai M.,2004,Maize mutants lacking chloroplast Fts Y exhibit pleiotropic defects in the biogenesis of thylakoid membranes,Plant Cell,16(1):201-214
Bae C.H.,Abe T.,Matsuyama T.,Fukunishi N.,Nagata N.,Nakano T.,Kaneko Y.,Miyoshi K.,Matsushima H.,and Yoshida,2001,Regulation of chloroplast gene expression is affected in ali,a novel tobacco albino mutant,Annals of Botany,88(4):545-553
Chang W.D.,Hui L.L.,Xu M.,Fu A.G.,and Wang Y.H.,2016,Rese arch progress of the plant chloroplast protease,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),35(3):728-739(常伟东,惠亮亮,徐敏,付爱根,王玉华,2016,植物叶绿体蛋白酶研究进展,基因组学与应用生物学,35(3):728-739)
Cheng H.,Li S.F.,Chen M.,Yu F.L.,Yan J.,Liu Y.M.,and Chen L.G.,1999,Physiological and biochemical essence of the extraordinary characters of Anji Baicha,Chaye Kexue(Journal of Tea Science),19(2):87-92(成浩,李素芳,陈明,虞富莲,晏静,刘益民,陈龙安,1999,安吉白茶特异性状的生理生化本质,茶叶科学,19(2):87-92)
Cheng J.F.,Chen G.Y.,and Shen Y.G.,2012,Relational analysis of leaf characteristics and photosynthetic capacities of plants,Zhongguo Shengtai Nongye Xuebao(Chinese Journal of Eco-Agriculture),20(4):466-473(程建峰,陈根云,沈允钢,2012,植物叶片特征与光合性能的关系,中国生态农业学报,20(4):466-473)
Du X.H.,Zhou X.J.,and Peng F.R.,2007,Comparison on net photosynthetic rate and tran sp iration rate of five tea cultivars,Linye Keji Kaifa(Journal of Forestry Engineering),21(4):21-24(杜旭华,周贤军,彭方仁,2007,不同茶树品种净光合与蒸腾速率比较,林业科技开发,21(4):21-24)
Gong Y.S.,Huang J.A.,Cui X.X.,Li Q.,Li Y.H.,and Chen J.H.,2008,Determination of theanine in tea by paired ion chromatograph,Chaye Kexue(Journal of Tea Science),28(2):89-92(龚雨顺,黄建安,崔湘兴,李勤,李银花,陈金华,2008,离子对色谱法测定茶叶中的茶氨酸,茶叶科学,28(2):89-92)
Jin J.,Luo Y.P.,Ren M.X.,and Kang M.L.,2003,Study on the photosynthet ic characteristics of graf ted tea plants,Chaye(Journal of Tea),29(2):86-88(金洁,骆耀平,任明兴,康孟利,2003,嫁接茶树光合特性研究,茶叶,29(2):86-88)
Jung K.H.,Hur J.,Ryu C.H.,Choi Y.,Chung Y.Y.,Miyao A.,Hirochika H.,and An G.,2003,Characterization of a rice chlorophyll-deficient mutant using the t-dna gene-trap system,Plant and Cell Physiology,44(5):463-472
Kito M.,Kokura H.,Izaki J.,and Sasaoka K.,1968,Theanine,a precursor of the phloroglucinol nucleus of catechins in tea plants,Phytochemistry,7(4):599-603
Li Q.,Huang J.,Liu S.,Li J.,Yang X.,Liu Y.,and Liu Z.,2011,Proteomic analysis of young leaves at three developmental stages in an albino tea cultivar,Proteome Science,9(1):44
Li S.F.,Chen S.R.,and Cheng H.,1995,Studies on the staging albescent phenom enon in the plants-Observation on the uitramicrostructure of the chlorplast,Chaye Kexue(Journal of Tea Science),15(1):23-26(李素芳,陈树尧,成浩,1995,茶树阶段性返白现象的研究-叶绿体超微结构的变化,茶叶科学,15(1):23-26)
Lin J.K.,Zhen J.G.,Yuan M.,Zhang X.Q.,and Wang F.,2005,Differential proteomic analysis in normal tea shoot and exogenous induced tea shoot for increasing EGCG content,Chaye Kexue(Journal of Tea Science),25(2):109-115(林金科,郑金贵,袁明,张学琴,王凤,2005,外源诱导提高茶树EGCG含量过程的蛋白质组差异分析,茶叶科学,25(2):109-115)
Lin Z.F.,Peng C.L.,Lin G.Z.,and Zhang J.L.,2003,Alteration of components of chlorophyll-protein complexes and distribution of excitation energy between the two photosystems in two new rice chlorophyll b-less mutants,Photosynthetica,41(4):589-595
Liu Y.L.,Liu F.,Zhou K.N.,Su X.M.,Fang X.W.,Zhang Y.H.,and Bao Y.Q.,2016,Phenotypic characterization and gene mapping of a thermo-sensitive albino leaf mutant tsa1 in rice,Zuowu Xuebao(Acta Agronomica Sinica),42(12):1754-1763(刘钰龙,刘峰,周坤能,苏晓妹,方先文,张云辉,鲍依群,2016,水稻温敏型叶片白化突变体tsa1的表型鉴定和基因定位,作物学报,42(12):1754-1763)
Luo P.G.,and Ren Z.L.,2006,Wheat leaf chlorosis controlled by a single recessive gene,Journal of Plant Physiology and Molecular Biology,32(3):330-338
Sawers R.J.,Viney J.,Farmer P.R.,Bussey R.R.,Olsefski G.,Anufrikova K.,Hunter C.N.,and Brutnell T.P.,2006,The maize Oil yellow1(Oy1)gene encodes the I subunit of magnesium chelatase,Plant Molecular Biology,60(1):95-106
Sun J.,Zhu L.G.,Lin Z.K.,and Zhang W.J.,2015,Research progress on photosynthesis of tea plant,Fujian Nongye Xuebao(Fujian Journal of Agricultural Sciences),30(12):1231-1237(孙君,朱留刚,林志坤,张文锦,2015,茶树光合作用研究进展,福建农业学报,30(12):1231-1237)
Sundberg E.,Slagter J.G.,Fridborg I.,Cleary S.P.,Robinson C.,and Coupland G.,1997,Albino3,an arabidopsis nuclear gene essential for chloroplast differentiation,encodes a chloroplast protein that shows homology to proteins present in bacterial membranes and yeast mitochondria,Plant Cell,9(5):717-730
Tan X.X.,and Tang Z.S.,1996,Leaf photosynthesis and chlorophyll fluorescence in a chlorophyll-deficient mutant of barley,Zhiwu Shengli Xuebao(Plant Physiology Journal),22(1):51-57(谭新星,汤泽生,1996,叶绿素缺乏的大麦突变体的光合作用和叶绿素荧光,植物生理学报,22(1):51-57)
Tang Q.,Ye S.R.,Shan H.L.,and Luo X.P.,2006,Comparative study on photosynthetic characteristics of the introducing tea cultivars,Sichuang Nongye Daxue Xuebao(Journal of Sichuan Agricultural University),24(3):303-308(唐茜,叶善蓉,单虹丽,罗学平,2006,引进茶树品种光合特性的比较研究,四川农业大学学报,24(3):303-308)
Wang Z.S.,Tong X.L.,and Zhu S.T.,1991,Quantitative analysis of catechins with HPLC,Chaye Kexue(Journal of Tea Science),11(S):93-99(王增盛,童小麟,朱尚同,1991,茶儿茶素的高效液相色谱测定方法,茶叶科学,11(S):93-99)
Wu D.X.,Shu Q.Y.,and Xia Y.W.,2002,In vitro mutagenesis induced novel thermo/photoperiod-sensitive genic male sterile indica rice with green-revertible xanthan leaf color marker,Euphytica,123(2):195-202
Wu L.Q.,You C.C.,Ke J.,and He Q.H.,2013,Chloroplast development and physiological characteristics of green-revertible albino leaf color mutants in rice,Redai Zuowu Xuebao(Chinese Journal of Tropical Crops),34(6):1115-1120(武立权,尤翠翠,柯建,何清华,2013,叶色白化水稻突变体转绿中若干生理与叶绿体发育特型的研究,热带作物学报,34(6):1115-1120)
Xiong L.,Li J.,Li Y.,Yuan L.,Liu S.,Huang J.,and Liu Z.,2013,Dynamic changes in catechin levels and catechin biosynthesis-related gene expression in albino tea plants(Camellia sinensis L.),Plant Physiology and Biochemistry,71:132-143
Xu H.R.,Tong Q.Q.,Luo Y.P.,Chen Y.Y.,and Hong J.,1992,Studies on photosynthetic characteristics of tea gremplasms from Zhejiang,Zhejiang Nongye Daxue Xuebao(Acta Agricultural University Zhejiangensis),18(S):48-52(须海荣,童启庆,骆耀平,陈玉银,洪健,1992,浙江茶树资源光合特性的研究,浙江农业大学学报,18(S):48-52)
Yuan Z.L.,Li H.X.,Lv L.Z.,and Han Y.,2010,Study on photosynthetic characteristics and quality components of introduced tea cultivars in southof Henan Province,Henan Nongye Daxue Xuebao(Journal of Henan Agricultural University),44(3):267-272(袁祖丽,李华鑫,吕立哲,韩莹,2010,豫南引种茶树品种光合特性与品质成分的研究,河南农业大学学报,44(3):267-272)
Zhang H.,Li J.,Yoo J.H.,Yoo S.C.,Cho S.H.,Koh H.J.,Seo H.S.,and Paek N.C.,2006,Rice chlorina-1 and chlorina-9 encode chld and chli subunits of mg-chelatase,a key enzyme for chlorophyll synthesis and chloroplast development,Plant Molecular Biology,62(3):325-337
Zhu M.K.,Hu Z.L.,Zhou S.,Li Y.L.,and Chen G.P.,2012,Research progress of plant leaf albino,Shengming Kexue(Chinese Bulletin of Life Sciences),24(3):255-261(朱明库,胡宗利,周爽,李亚丽,陈国平,2012,植物叶色白化研究进展,生命科学,24(3):255-261)
Asakura Y.,Hi rohashi T.,Kikuchi S.,Belcher S.,Osborne E.,Yano S.,Terashima I.,Barkan A.,and Nakai M.,2004,Maize mutants lacking chloroplast Fts Y exhibit pleiotropic defects in the biogenesis of thylakoid membranes,Plant Cell,16(1):201-214
Bae C.H.,Abe T.,Matsuyama T.,Fukunishi N.,Nagata N.,Nakano T.,Kaneko Y.,Miyoshi K.,Matsushima H.,and Yoshida,2001,Regulation of chloroplast gene expression is affected in ali,a novel tobacco albino mutant,Annals of Botany,88(4):545-553
Chang W.D.,Hui L.L.,Xu M.,Fu A.G.,and Wang Y.H.,2016,Rese arch progress of the plant chloroplast protease,Jiyinzuxue Yu Yingyong Shengwuxue(Genomics and Applied Biology),35(3):728-739(常伟东,惠亮亮,徐敏,付爱根,王玉华,2016,植物叶绿体蛋白酶研究进展,基因组学与应用生物学,35(3):728-739)
Cheng H.,Li S.F.,Chen M.,Yu F.L.,Yan J.,Liu Y.M.,and Chen L.G.,1999,Physiological and biochemical essence of the extraordinary characters of Anji Baicha,Chaye Kexue(Journal of Tea Science),19(2):87-92(成浩,李素芳,陈明,虞富莲,晏静,刘益民,陈龙安,1999,安吉白茶特异性状的生理生化本质,茶叶科学,19(2):87-92)
Cheng J.F.,Chen G.Y.,and Shen Y.G.,2012,Relational analysis of leaf characteristics and photosynthetic capacities of plants,Zhongguo Shengtai Nongye Xuebao(Chinese Journal of Eco-Agriculture),20(4):466-473(程建峰,陈根云,沈允钢,2012,植物叶片特征与光合性能的关系,中国生态农业学报,20(4):466-473)
Du X.H.,Zhou X.J.,and Peng F.R.,2007,Comparison on net photosynthetic rate and tran sp iration rate of five tea cultivars,Linye Keji Kaifa(Journal of Forestry Engineering),21(4):21-24(杜旭华,周贤军,彭方仁,2007,不同茶树品种净光合与蒸腾速率比较,林业科技开发,21(4):21-24)
Gong Y.S.,Huang J.A.,Cui X.X.,Li Q.,Li Y.H.,and Chen J.H.,2008,Determination of theanine in tea by paired ion chromatograph,Chaye Kexue(Journal of Tea Science),28(2):89-92(龚雨顺,黄建安,崔湘兴,李勤,李银花,陈金华,2008,离子对色谱法测定茶叶中的茶氨酸,茶叶科学,28(2):89-92)
Jin J.,Luo Y.P.,Ren M.X.,and Kang M.L.,2003,Study on the photosynthet ic characteristics of graf ted tea plants,Chaye(Journal of Tea),29(2):86-88(金洁,骆耀平,任明兴,康孟利,2003,嫁接茶树光合特性研究,茶叶,29(2):86-88)
Jung K.H.,Hur J.,Ryu C.H.,Choi Y.,Chung Y.Y.,Miyao A.,Hirochika H.,and An G.,2003,Characterization of a rice chlorophyll-deficient mutant using the t-dna gene-trap system,Plant and Cell Physiology,44(5):463-472
Kito M.,Kokura H.,Izaki J.,and Sasaoka K.,1968,Theanine,a precursor of the phloroglucinol nucleus of catechins in tea plants,Phytochemistry,7(4):599-603
Li Q.,Huang J.,Liu S.,Li J.,Yang X.,Liu Y.,and Liu Z.,2011,Proteomic analysis of young leaves at three developmental stages in an albino tea cultivar,Proteome Science,9(1):44
Li S.F.,Chen S.R.,and Cheng H.,1995,Studies on the staging albescent phenom enon in the plants-Observation on the uitramicrostructure of the chlorplast,Chaye Kexue(Journal of Tea Science),15(1):23-26(李素芳,陈树尧,成浩,1995,茶树阶段性返白现象的研究-叶绿体超微结构的变化,茶叶科学,15(1):23-26)
Lin J.K.,Zhen J.G.,Yuan M.,Zhang X.Q.,and Wang F.,2005,Differential proteomic analysis in normal tea shoot and exogenous induced tea shoot for increasing EGCG content,Chaye Kexue(Journal of Tea Science),25(2):109-115(林金科,郑金贵,袁明,张学琴,王凤,2005,外源诱导提高茶树EGCG含量过程的蛋白质组差异分析,茶叶科学,25(2):109-115)
Lin Z.F.,Peng C.L.,Lin G.Z.,and Zhang J.L.,2003,Alteration of components of chlorophyll-protein complexes and distribution of excitation energy between the two photosystems in two new rice chlorophyll b-less mutants,Photosynthetica,41(4):589-595
Liu Y.L.,Liu F.,Zhou K.N.,Su X.M.,Fang X.W.,Zhang Y.H.,and Bao Y.Q.,2016,Phenotypic characterization and gene mapping of a thermo-sensitive albino leaf mutant tsa1 in rice,Zuowu Xuebao(Acta Agronomica Sinica),42(12):1754-1763(刘钰龙,刘峰,周坤能,苏晓妹,方先文,张云辉,鲍依群,2016,水稻温敏型叶片白化突变体tsa1的表型鉴定和基因定位,作物学报,42(12):1754-1763)
Luo P.G.,and Ren Z.L.,2006,Wheat leaf chlorosis controlled by a single recessive gene,Journal of Plant Physiology and Molecular Biology,32(3):330-338
Sawers R.J.,Viney J.,Farmer P.R.,Bussey R.R.,Olsefski G.,Anufrikova K.,Hunter C.N.,and Brutnell T.P.,2006,The maize Oil yellow1(Oy1)gene encodes the I subunit of magnesium chelatase,Plant Molecular Biology,60(1):95-106
Sun J.,Zhu L.G.,Lin Z.K.,and Zhang W.J.,2015,Research progress on photosynthesis of tea plant,Fujian Nongye Xuebao(Fujian Journal of Agricultural Sciences),30(12):1231-1237(孙君,朱留刚,林志坤,张文锦,2015,茶树光合作用研究进展,福建农业学报,30(12):1231-1237)
Sundberg E.,Slagter J.G.,Fridborg I.,Cleary S.P.,Robinson C.,and Coupland G.,1997,Albino3,an arabidopsis nuclear gene essential for chloroplast differentiation,encodes a chloroplast protein that shows homology to proteins present in bacterial membranes and yeast mitochondria,Plant Cell,9(5):717-730
Tan X.X.,and Tang Z.S.,1996,Leaf photosynthesis and chlorophyll fluorescence in a chlorophyll-deficient mutant of barley,Zhiwu Shengli Xuebao(Plant Physiology Journal),22(1):51-57(谭新星,汤泽生,1996,叶绿素缺乏的大麦突变体的光合作用和叶绿素荧光,植物生理学报,22(1):51-57)
Tang Q.,Ye S.R.,Shan H.L.,and Luo X.P.,2006,Comparative study on photosynthetic characteristics of the introducing tea cultivars,Sichuang Nongye Daxue Xuebao(Journal of Sichuan Agricultural University),24(3):303-308(唐茜,叶善蓉,单虹丽,罗学平,2006,引进茶树品种光合特性的比较研究,四川农业大学学报,24(3):303-308)
Wang Z.S.,Tong X.L.,and Zhu S.T.,1991,Quantitative analysis of catechins with HPLC,Chaye Kexue(Journal of Tea Science),11(S):93-99(王增盛,童小麟,朱尚同,1991,茶儿茶素的高效液相色谱测定方法,茶叶科学,11(S):93-99)
Wu D.X.,Shu Q.Y.,and Xia Y.W.,2002,In vitro mutagenesis induced novel thermo/photoperiod-sensitive genic male sterile indica rice with green-revertible xanthan leaf color marker,Euphytica,123(2):195-202
Wu L.Q.,You C.C.,Ke J.,and He Q.H.,2013,Chloroplast development and physiological characteristics of green-revertible albino leaf color mutants in rice,Redai Zuowu Xuebao(Chinese Journal of Tropical Crops),34(6):1115-1120(武立权,尤翠翠,柯建,何清华,2013,叶色白化水稻突变体转绿中若干生理与叶绿体发育特型的研究,热带作物学报,34(6):1115-1120)
Xiong L.,Li J.,Li Y.,Yuan L.,Liu S.,Huang J.,and Liu Z.,2013,Dynamic changes in catechin levels and catechin biosynthesis-related gene expression in albino tea plants(Camellia sinensis L.),Plant Physiology and Biochemistry,71:132-143
Xu H.R.,Tong Q.Q.,Luo Y.P.,Chen Y.Y.,and Hong J.,1992,Studies on photosynthetic characteristics of tea gremplasms from Zhejiang,Zhejiang Nongye Daxue Xuebao(Acta Agricultural University Zhejiangensis),18(S):48-52(须海荣,童启庆,骆耀平,陈玉银,洪健,1992,浙江茶树资源光合特性的研究,浙江农业大学学报,18(S):48-52)
Yuan Z.L.,Li H.X.,Lv L.Z.,and Han Y.,2010,Study on photosynthetic characteristics and quality components of introduced tea cultivars in southof Henan Province,Henan Nongye Daxue Xuebao(Journal of Henan Agricultural University),44(3):267-272(袁祖丽,李华鑫,吕立哲,韩莹,2010,豫南引种茶树品种光合特性与品质成分的研究,河南农业大学学报,44(3):267-272)
Zhang H.,Li J.,Yoo J.H.,Yoo S.C.,Cho S.H.,Koh H.J.,Seo H.S.,and Paek N.C.,2006,Rice chlorina-1 and chlorina-9 encode chld and chli subunits of mg-chelatase,a key enzyme for chlorophyll synthesis and chloroplast development,Plant Molecular Biology,62(3):325-337
Zhu M.K.,Hu Z.L.,Zhou S.,Li Y.L.,and Chen G.P.,2012,Research progress of plant leaf albino,Shengming Kexue(Chinese Bulletin of Life Sciences),24(3):255-261(朱明库,胡宗利,周爽,李亚丽,陈国平,2012,植物叶色白化研究进展,生命科学,24(3):255-261)