一个新的黄瓜叶色突变体的光合特性分析
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摘要
叶色突变体是研究植物光合系统结构与功能、叶绿素生物合成与代谢以及叶绿体发育的理想材料。该研究以黄瓜黄绿叶突变体C777及其野生型CCMC为试材,对其苗期叶片的光合色素质量分数、光合特性、叶绿素荧光参数等进行测定。结果表明,突变体叶绿素a、叶绿素b、类胡萝卜素以及总叶绿素质量分数均显著低于野生型;光合特性方面,突变体净光合速率显著低于野生型,而胞间CO_2浓度显著高于野生型,表明突变体对CO2的利用率较野生型低;叶绿素荧光动力学参数的测定结果表明,突变体的F_o、F_m、F_v/F_m、F′_v/F′_m、Φ_(PSⅡ)、qP、ETR和NPQ均显著降低,说明光合色素质量分数的降低导致PSⅡ反应中心捕光能力较弱和光化学转化效率较低,使得突变体光合速率降低。试验为进一步阐明该黄瓜黄绿叶突变体的叶色突变机制和该种质的合理利用提供相关基础资料。
Leaf color mutant is an ideal material for studying photosynthesis system structure and function,chlorophyll biosynthesis and metabolism,and chloroplast development.The mutant C777 and wild type CCMC were taken as materials.The photosynthetic pigment mass fraction,photosynthetic characteristics,chlorophyll fluorescence parameters were measured in this study.The results showed that chlorophyll a,chlorophyll b,carotenoid and total chlorophyll mass fraction of mutant were significantly lower than those of wild type.In photosynthetic characteristics,the net photosynthetic rate of the mutant was significantly lower than that of the wild type,while the intercellular CO_2 concentration was significantly higher than that of the wild type,it indicated that the utilization of CO_2 by mutant was lower than wild type.The determination of chlorophyll fluorescence kinetic revealed that the mutant significantly decreased in F_o,F_m,F_v/F_m,F′_v/F′_m,Φ_(PSⅡ),qP,ETR and NPQ,this suggested that the reduction in photosynthetic pigment mass fraction led to low PSⅡreaction,light harvest ability,and less photochemical conversion efficiency,so the mutant photosynthetic rate decreased.These results provide basic information for further elucidating mechanism of the leaf color mutation and rational utilization of the germplasm in the virescent leaf mutant in cucumber.
Leaf color mutant is an ideal material for studying photosynthesis system structure and function,chlorophyll biosynthesis and metabolism,and chloroplast development.The mutant C777 and wild type CCMC were taken as materials.The photosynthetic pigment mass fraction,photosynthetic characteristics,chlorophyll fluorescence parameters were measured in this study.The results showed that chlorophyll a,chlorophyll b,carotenoid and total chlorophyll mass fraction of mutant were significantly lower than those of wild type.In photosynthetic characteristics,the net photosynthetic rate of the mutant was significantly lower than that of the wild type,while the intercellular CO_2 concentration was significantly higher than that of the wild type,it indicated that the utilization of CO_2 by mutant was lower than wild type.The determination of chlorophyll fluorescence kinetic revealed that the mutant significantly decreased in F_o,F_m,F_v/F_m,F′_v/F′_m,Φ_(PSⅡ),qP,ETR and NPQ,this suggested that the reduction in photosynthetic pigment mass fraction led to low PSⅡreaction,light harvest ability,and less photochemical conversion efficiency,so the mutant photosynthetic rate decreased.These results provide basic information for further elucidating mechanism of the leaf color mutation and rational utilization of the germplasm in the virescent leaf mutant in cucumber.
引文
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[7]刘国民,高必军,文绍山,等.具有黄绿叶标记的香型籼稻不育系选育研究[J].中国农业科学,2010,43(4):855-861.LIU G M,GAO B J,WEN SH SH,et al.Breeding of male sterile line of aromatic indica rice with yellow-green leaves marker[J].Scientia Agricultura Sinica,2010,43(4):855-861.
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[9]LI R Q,HUANG J Z,ZHAO H J,et al.A down-regulated epi-allele of the genomes uncoupled 4gene generates a xantha marker trait in rice[J].Theoretical and Applied Genetics,2014,127(11):2491-2501.
[10]田明爽,宋美珍,范术丽,等.植物叶绿素缺失突变体分子机制研究进展[J].西北植物学报,2011,31(9):1900-1907.TIAN M SH,SONG M ZH,FAN SH L,et al.Advance in research of molecular mechanism of chlorophyll-deficient mutants in plants[J].Acta Botanica Boreali-Occidentalia Sinica,2011,31(9):1900-1907.
[11]李秀兰,孙小秋,王平荣,等.一个新的水稻黄绿叶突变体的遗传分析与基因定位[J].作物学报,2010,36(6):1050-1054.LI X L,SUN X Q,WANG P R,et al.Genetic analysis and gene mapping of a novel yellow-green leaf mutant in rice[J].Acta Agronomica Sinica,2010,36(6):1050-1054.
[12]REED S,ATKINSON T,GORECKI C,et al.Candidate gene identification for a lethal chlorophyll-deficient mutant in soybean[J].Agronomy,2014,4(4):462-469.
[13]GUAN H Y,XU X B,HE C M,et al.Fine mapping and candidate gene analysis of the leaf-color gene ygl-1in maize[J].PLoS One,2016,11(4):e0153962.
[14]MA X Z,SUN X Q,LI C M,et al.Map-based cloning and characterization of the novel yellow-green leaf gene ys83in rice(Oryza sativa)[J].Plant Physiology&Biochemistry,2017,111:1-9.
[15]DENG X J,ZHANG H Q,WANG Y,et al.Mapped clone and functional analysis of leaf-color gene ygl7in a rice hybrid(Oryza sativa L.spp.indica)[J].PLoS One,2014,9(6):e99564.
[16]苗晗,顾兴芳,张圣平,等.黄瓜黄绿叶突变体光合色素变化及相关基因差异表达[J].中国农业科学,2010,43(19):4027-4035.MIAO H,GU X F,ZHANG SH P,et al.Changes of the photosynthetic pigment and differential expression of the correlated genes in a chlorophyll-deficient cucumber mutant(Cucumis sativus L.)[J].Scientia Agricultura Sinica,2010,43(19):4027-4035.
[17]PIERCE L K,WEHNER T C.Review of genes and linkage groups in cucumber[J].Hortscience,1990,25(6):605-615.
[18]WALTERS S A,SHETTY N V,WEHNER T C.Segregation and linkage of several genes in cucumber[J].Journal of the American Society for Horticultural Science,2001,126(4):442-450.
[19]MIAO H,ZHANG S P,WANG M,et al.Fine mapping of virescent leaf gene v-1in cucumber(Cucumis sativus L.)[J].International Journal of Molecular Sciences,2016,17(10):1602.
[20]GAO M L,HU L L,LI Y H,et al.The chlorophyll-deficient golden leaf mutation in cucumber is due to a single nucleotide substitution in CsChlIfor magnesium chelataseⅠsubunit[J].Theoretical and Applied Genetics,2016,129(10):1961-1973.
[21]李万青,高波,杨俊,等.一个新的黄瓜叶色黄化突变体的生理特性分析[J].西北农业学报,2015,24(7):98-103.LI W Q,GAO B,YANG J,et al.Physiological characteristic analysis of a new leaf color yellow mutant in cucumber[J].Acta Agriculturae Boreali-occidentalis Sinica,2015,24(7):98-103.
[22]HILL C M,PEARSON S A,SMITH A J,et al.Inhibition of chlorophyll synthesis in Hordeum vulgare by 3-amino2,3-dihydrobenzoic acid(gabaculin)[J].Bioscience Reports,1985,5(9):775-781.
[23]于凤,高丽,闫志坚,等.库布齐沙地6种植物叶绿素荧光参数比较[J].草业科学,2012,29(1):83-87.YU F,GAO L,YAN ZH J,et al.Differentiation of chlorophyll fluorescence parameters of six plants species in the Kubuqi sandy land[J].Pratacultural Science,2012,29(1):83-87.
[24]ROHEK K,BARTK M.Technique of the modulated chlorophyll fluorescence:basic concepts,useful parameters,and some applications[J].Photosynthetica,1999,37(3):339-363.
[25]王家训,苏晓东.黄瓜黄绿叶突变性状的遗传分析[J].遗传,2000,22(5):313-315.WANG J X,SU X D.Genetic analysis of a yellow green leaf mutant character in cucumber(Cucumis sativus L.)[J].Hereditas,2000,22(5):313-315.
[26]常青山,陈发棣,腾年军,等.菊花黄绿叶突变体不同类型叶片的叶绿素含量和结构特征比较[J].西北植物学报,2008,28(9):1772-1777.CHANG Q SH,CHEN F D,TENG N J,et al.Chlorophyll mass fractions and chloroplast microstructure and ultrastructure of leaves in yellow-green mutant of chrysanthemum[J].Acta Botanica Boreali-Occidentalia Sinica,2008,28(9):1772-1777.
[27]杨佳秀,杜丽芬,刘录祥,等.小麦旗叶黄化转绿突变体的生理分析及细胞学研究[J].西北植物学报,2015,35(12):2455-2461.YANG J X,DU L F,LIU L X,et al.Physiological analysis and cytological observation of the wheat mutant with yellow green-revertible flag leaf[J].Acta Botanica Boreali-Occidentalia Sinica,2015,35(12):2455-2461.
[28]李晓,冯伟,曾晓春.叶绿素荧光分析技术及应用进展[J].西北植物学报,2006,26(10):2186-2196.LI X,FENG W,ZENG X CH.Advances in chlorophyll fluorescence analysis and its uses[J].Acta Botanica BorealiOccidentalia Sinica,2006,26(10):2186-2196.
[29]曹莉,王辉,孙道杰,等.小麦黄化突变体光合作用及叶绿素荧光特性研究[J].西北植物学报,2006,26(10):2083-2087.CAO L,WANG H,SUN D J,et al.Photosynthesis and chlorophyll fluorescence characters of xantha wheat mutant[J].Acta Botanica Boreali-Occidentalia Sinica,2006,26(10):2083-2087.
[30]常青山,张利霞,陈煜,等.菊花黄绿叶突变体的叶绿素荧光特性[J].江苏农业学报,2011,27(5):1093-1099.CHANG Q SH,ZHANG L X,CHEN Y,et al.Chlorophyll fluorescence characteristics in yellow green leaf mutant of chrysanthemum[J].Jiangsu Journal of Agricultural Sciences,2011,27(5):1093-1099.
[31]欧立军.水稻叶色突变体的高光合特性[J].作物学报,2011,37(10):1860-1867.OU L J.High photosynthetic efficiency of leaf colour mutant of rice(Oryza sativa L.)[J].Acta Agronomica Sinica,2011,37(10):1860-1867.
[32]郭士伟,张云华,金永庆,等.小白菜(Brassica chinensis L.)黄苗突变体的叶绿素荧光特性[J].作物学报,2003,29(6):958-960.GUO SH W,ZHANG Y H,JIN Y Q,et al.Characterization of chlorophyll fluorescence in a mutant of Brassica chinensis L.with xantha seedling leaves[J].Acta Agronomica Sinica,2003,29(6):958-960.
[33]ZHONG X M,SUN S F,LI F H,et al.Photosynthesis of a yellow-green mutant line in maize[J].Photosynthetica,2015,53(4):499-505.
[2]LIU X P,YANG C,HAN F Q,et al.Genetics and fine mapping of a yellow-green leaf gene(ygl-1)in cabbage(Brassica oleracea var.capitata L.)[J].Molecular Breeding,2016,36(6):82.
[3]FAMBRINI M,CASTAGNA A,VECCHIA F D,et al.Characterization of a pigment-deficient mutant of sunflower(Helianthus annuus L.)with abnormal cloroplast biogenesis,reduced PSⅡactivity and low endogenous level of abscisic acid[J].Plant Science,2004,167(1):79-89.
[4]RUAN B P,GAO Z Y,ZHAO J,et al.The rice YGL gene encoding an Mg2+-chelatase ChlD subunit is affected by temperature for chlorophyll biosynthesis[J].Journal of Plant Biology,2017,60(4):314-321.
[5]ZHANG L L,LIU C,AN X Y,et al.Identification and genetic mapping of a novel incompletely dominant yellow leaf color gene,Y1718,on chromosome 2BS in wheat[J].Euphytica,2017,213(7):141.
[6]苗晗,顾兴芳,张圣平,等.蔬菜叶色突变体研究进展[J].中国蔬菜,2007,1(6):39-42.MIAO H,GU X F,ZHANG SH P,et al.Review on leaf color mutant of vegetable[J].China Vegetables,2007,1(6):39-42.
[7]刘国民,高必军,文绍山,等.具有黄绿叶标记的香型籼稻不育系选育研究[J].中国农业科学,2010,43(4):855-861.LIU G M,GAO B J,WEN SH SH,et al.Breeding of male sterile line of aromatic indica rice with yellow-green leaves marker[J].Scientia Agricultura Sinica,2010,43(4):855-861.
[8]ZHOU X S,SHEN S Q,WU D X,et al.Introduction of a xantha mutation for testing and increasing varietal purity in hybrid rice[J].Field Crops Research,2006,96(1):71-79.
[9]LI R Q,HUANG J Z,ZHAO H J,et al.A down-regulated epi-allele of the genomes uncoupled 4gene generates a xantha marker trait in rice[J].Theoretical and Applied Genetics,2014,127(11):2491-2501.
[10]田明爽,宋美珍,范术丽,等.植物叶绿素缺失突变体分子机制研究进展[J].西北植物学报,2011,31(9):1900-1907.TIAN M SH,SONG M ZH,FAN SH L,et al.Advance in research of molecular mechanism of chlorophyll-deficient mutants in plants[J].Acta Botanica Boreali-Occidentalia Sinica,2011,31(9):1900-1907.
[11]李秀兰,孙小秋,王平荣,等.一个新的水稻黄绿叶突变体的遗传分析与基因定位[J].作物学报,2010,36(6):1050-1054.LI X L,SUN X Q,WANG P R,et al.Genetic analysis and gene mapping of a novel yellow-green leaf mutant in rice[J].Acta Agronomica Sinica,2010,36(6):1050-1054.
[12]REED S,ATKINSON T,GORECKI C,et al.Candidate gene identification for a lethal chlorophyll-deficient mutant in soybean[J].Agronomy,2014,4(4):462-469.
[13]GUAN H Y,XU X B,HE C M,et al.Fine mapping and candidate gene analysis of the leaf-color gene ygl-1in maize[J].PLoS One,2016,11(4):e0153962.
[14]MA X Z,SUN X Q,LI C M,et al.Map-based cloning and characterization of the novel yellow-green leaf gene ys83in rice(Oryza sativa)[J].Plant Physiology&Biochemistry,2017,111:1-9.
[15]DENG X J,ZHANG H Q,WANG Y,et al.Mapped clone and functional analysis of leaf-color gene ygl7in a rice hybrid(Oryza sativa L.spp.indica)[J].PLoS One,2014,9(6):e99564.
[16]苗晗,顾兴芳,张圣平,等.黄瓜黄绿叶突变体光合色素变化及相关基因差异表达[J].中国农业科学,2010,43(19):4027-4035.MIAO H,GU X F,ZHANG SH P,et al.Changes of the photosynthetic pigment and differential expression of the correlated genes in a chlorophyll-deficient cucumber mutant(Cucumis sativus L.)[J].Scientia Agricultura Sinica,2010,43(19):4027-4035.
[17]PIERCE L K,WEHNER T C.Review of genes and linkage groups in cucumber[J].Hortscience,1990,25(6):605-615.
[18]WALTERS S A,SHETTY N V,WEHNER T C.Segregation and linkage of several genes in cucumber[J].Journal of the American Society for Horticultural Science,2001,126(4):442-450.
[19]MIAO H,ZHANG S P,WANG M,et al.Fine mapping of virescent leaf gene v-1in cucumber(Cucumis sativus L.)[J].International Journal of Molecular Sciences,2016,17(10):1602.
[20]GAO M L,HU L L,LI Y H,et al.The chlorophyll-deficient golden leaf mutation in cucumber is due to a single nucleotide substitution in CsChlIfor magnesium chelataseⅠsubunit[J].Theoretical and Applied Genetics,2016,129(10):1961-1973.
[21]李万青,高波,杨俊,等.一个新的黄瓜叶色黄化突变体的生理特性分析[J].西北农业学报,2015,24(7):98-103.LI W Q,GAO B,YANG J,et al.Physiological characteristic analysis of a new leaf color yellow mutant in cucumber[J].Acta Agriculturae Boreali-occidentalis Sinica,2015,24(7):98-103.
[22]HILL C M,PEARSON S A,SMITH A J,et al.Inhibition of chlorophyll synthesis in Hordeum vulgare by 3-amino2,3-dihydrobenzoic acid(gabaculin)[J].Bioscience Reports,1985,5(9):775-781.
[23]于凤,高丽,闫志坚,等.库布齐沙地6种植物叶绿素荧光参数比较[J].草业科学,2012,29(1):83-87.YU F,GAO L,YAN ZH J,et al.Differentiation of chlorophyll fluorescence parameters of six plants species in the Kubuqi sandy land[J].Pratacultural Science,2012,29(1):83-87.
[24]ROHEK K,BARTK M.Technique of the modulated chlorophyll fluorescence:basic concepts,useful parameters,and some applications[J].Photosynthetica,1999,37(3):339-363.
[25]王家训,苏晓东.黄瓜黄绿叶突变性状的遗传分析[J].遗传,2000,22(5):313-315.WANG J X,SU X D.Genetic analysis of a yellow green leaf mutant character in cucumber(Cucumis sativus L.)[J].Hereditas,2000,22(5):313-315.
[26]常青山,陈发棣,腾年军,等.菊花黄绿叶突变体不同类型叶片的叶绿素含量和结构特征比较[J].西北植物学报,2008,28(9):1772-1777.CHANG Q SH,CHEN F D,TENG N J,et al.Chlorophyll mass fractions and chloroplast microstructure and ultrastructure of leaves in yellow-green mutant of chrysanthemum[J].Acta Botanica Boreali-Occidentalia Sinica,2008,28(9):1772-1777.
[27]杨佳秀,杜丽芬,刘录祥,等.小麦旗叶黄化转绿突变体的生理分析及细胞学研究[J].西北植物学报,2015,35(12):2455-2461.YANG J X,DU L F,LIU L X,et al.Physiological analysis and cytological observation of the wheat mutant with yellow green-revertible flag leaf[J].Acta Botanica Boreali-Occidentalia Sinica,2015,35(12):2455-2461.
[28]李晓,冯伟,曾晓春.叶绿素荧光分析技术及应用进展[J].西北植物学报,2006,26(10):2186-2196.LI X,FENG W,ZENG X CH.Advances in chlorophyll fluorescence analysis and its uses[J].Acta Botanica BorealiOccidentalia Sinica,2006,26(10):2186-2196.
[29]曹莉,王辉,孙道杰,等.小麦黄化突变体光合作用及叶绿素荧光特性研究[J].西北植物学报,2006,26(10):2083-2087.CAO L,WANG H,SUN D J,et al.Photosynthesis and chlorophyll fluorescence characters of xantha wheat mutant[J].Acta Botanica Boreali-Occidentalia Sinica,2006,26(10):2083-2087.
[30]常青山,张利霞,陈煜,等.菊花黄绿叶突变体的叶绿素荧光特性[J].江苏农业学报,2011,27(5):1093-1099.CHANG Q SH,ZHANG L X,CHEN Y,et al.Chlorophyll fluorescence characteristics in yellow green leaf mutant of chrysanthemum[J].Jiangsu Journal of Agricultural Sciences,2011,27(5):1093-1099.
[31]欧立军.水稻叶色突变体的高光合特性[J].作物学报,2011,37(10):1860-1867.OU L J.High photosynthetic efficiency of leaf colour mutant of rice(Oryza sativa L.)[J].Acta Agronomica Sinica,2011,37(10):1860-1867.
[32]郭士伟,张云华,金永庆,等.小白菜(Brassica chinensis L.)黄苗突变体的叶绿素荧光特性[J].作物学报,2003,29(6):958-960.GUO SH W,ZHANG Y H,JIN Y Q,et al.Characterization of chlorophyll fluorescence in a mutant of Brassica chinensis L.with xantha seedling leaves[J].Acta Agronomica Sinica,2003,29(6):958-960.
[33]ZHONG X M,SUN S F,LI F H,et al.Photosynthesis of a yellow-green mutant line in maize[J].Photosynthetica,2015,53(4):499-505.