缺铁胁迫柑橘砧木幼苗的光合特性和叶绿体超微结构
|
摘要
【目的】通过研究枳壳与枳橙砧木在缺铁和正常铁浓度处理下的反应,重点揭示两种柑橘砧木光合特性、叶绿体超微结构等对铁敏感性的差异。【方法】以柑橘的枳壳砧木和枳橙砧木实生苗为试验材料,设置缺铁(–Fe,0μmol/L)和正常铁(+Fe,37.3μmol/L)2个处理进行营养液培养,测定了缺铁胁迫对两种砧木苗期铁元素含量与积累量、光合色素含量、叶片糖含量的影响,并进行了叶绿体超微结构的电镜扫描。【结果】缺铁胁迫显著降低了两种砧木铁元素含量与积累量、叶片光合色素含量,且枳橙砧木光合色素含量下降幅度较大。与对照相比,缺铁后枳壳砧木叶片的糖类物质含量降低,且达到显著差异水平;而枳橙砧木叶片可溶性糖、蔗糖含量显著升高,淀粉和果糖含量显著下降。另外,缺铁胁迫条件下,两种柑橘砧木片层结构模糊,嗜饿体数目增加。并且,缺铁后枳壳砧木叶绿体长度、厚度比对照分别降低了22.1%、26.4%,枳橙砧木则分别下降了55.1%、40.4%。【结论】缺铁胁迫下,枳橙砧木幼苗的铁元素含量和积累量、叶片光合色素含量、叶绿体超微结构等均比枳壳砧木受到较大影响,表明枳橙砧木比枳壳砧木对铁营养缺乏更加敏感。
【Objectives】Response of photosynthetic characteristics and chloroplast ultrastructure of different Fe sensitivity citrus rootstocks to different iron(Fe) treatments was studied.【Methods】A hydroponic experiment was carried out with –Fe(0 μmol/L) and +Fe(37.3 μmol/L), and citrange and trifoliate orange rootstocks were selected as research materials. The contents of Fe, photosynthetic pigment and carbohydrates and chloroplast ultrastructure of samples were analyzed.【Results】The results showed that Fe deficiency significantly decreased the contents and accumulation of Fe, photosynthetic pigment contents in both rootstocks, and the photosynthetic pigment content in citrange rootstock was decreased in a larger extent. The iron deficiency significantly decreased carbohydrates contents in leaves of trifoliate orange, and the contents of total soluble sugar and glucose in leaves of citrange were significantly increased under Fe-deficiency, however, the contents of fructose and starch in leaves of citrange were significantly decreased with no Fe supply. Lamellae structure in chloroplast became fuzzy, and the numbers of plastoglobulus were increased without the Fe application.Compared with the Fe-sufficiency treatment,the chloroplast length and the chloroplast thickness of trifoliate were decreased by 22.1% and 26.4%, and those of the citrange orange were decreased by 55.1% and 40.4%.【Conclusions】The content and accumulation of iron, the content of photosynthetic pigment and the ultrastructure of chloroplast of citrange rootstocks were more sensitive to Fe deficiency.
【Objectives】Response of photosynthetic characteristics and chloroplast ultrastructure of different Fe sensitivity citrus rootstocks to different iron(Fe) treatments was studied.【Methods】A hydroponic experiment was carried out with –Fe(0 μmol/L) and +Fe(37.3 μmol/L), and citrange and trifoliate orange rootstocks were selected as research materials. The contents of Fe, photosynthetic pigment and carbohydrates and chloroplast ultrastructure of samples were analyzed.【Results】The results showed that Fe deficiency significantly decreased the contents and accumulation of Fe, photosynthetic pigment contents in both rootstocks, and the photosynthetic pigment content in citrange rootstock was decreased in a larger extent. The iron deficiency significantly decreased carbohydrates contents in leaves of trifoliate orange, and the contents of total soluble sugar and glucose in leaves of citrange were significantly increased under Fe-deficiency, however, the contents of fructose and starch in leaves of citrange were significantly decreased with no Fe supply. Lamellae structure in chloroplast became fuzzy, and the numbers of plastoglobulus were increased without the Fe application.Compared with the Fe-sufficiency treatment,the chloroplast length and the chloroplast thickness of trifoliate were decreased by 22.1% and 26.4%, and those of the citrange orange were decreased by 55.1% and 40.4%.【Conclusions】The content and accumulation of iron, the content of photosynthetic pigment and the ultrastructure of chloroplast of citrange rootstocks were more sensitive to Fe deficiency.
引文
[1]Castle W,Tucker D,Krezdorn A,Youtsey C.Rootstocks for Florida citrus[M].Gainesville:University of Florida,1993:1-22.
[2]周开兵,郭文武,夏仁学,等.两种柑桔体细胞杂种砧木利用价值和砧穗互作生化机制的探讨[J].园艺学报,2004,31(4):427-432.Zhou K B,Guo W W,Xia R X,et al.Studies on the utilization potential of two somatic hybrid rootstocks and the biochemical mechanism of rootstock-scion interaction[J].Acta Horticulturae Sinica,2004,31(4):427-432.
[3]Sheng O,Song S,Peng S,et al.The effects of low boron on growth,gas exchange,boron concentration and distribution of‘Newhall’navel orange(Citrus sinensis,Osb.)plants grafted on two rootstocks[J].Scientia Horticulturae,2009,121(3):278-283.
[4]Hellín E,Hernández-Cortés J A,Piqueras A,et al.The influence of the iron content on the superoxide dismutase activity and chloroplast ultrastructure of Citrus limon[A].Iron nutrition in soils and plants[M].Springer Netherlands,1995:211-229.
[5]姚允聪,王绍辉,孔云.弱光条件下桃叶片结构及光合特性与叶绿体超微结构变化[J].中国农业科学,2007,40(4):855-863.Yao Y C,Wang S H,Kong Y.Characteristics of photosynthesis mechanism in different peach species under low light intensity[J].Scientia Agricultura Sinica,2007,40(4):855-863.
[6]郭卫东,桑丹,郑建树,等.缺氮对佛手气体交换、叶绿素荧光及叶绿体超微结构的影响[J].浙江大学学报(农业与生命科学版),2009,35(3):307-314.Guo W D,Sang D,Zheng S J,et al.Effects of nitrogen deficiency on the gas exchange,chlorophyll fluorescence and chloroplast ultrastructure in fingered citron[J].Journal of Zhejiang University(Agricultural&Life Sciences),2009,35(3):307-314.
[7]王跃华,张丽霞,孙其远.钙过量对茶树光合特性及叶绿体超微结构的影响[J].植物营养与肥料学报,2010,16(2):432-438.Wang Y H,Zhang L X,Sun Q Y.Effects of excessive calcium fertilization on photosynthetic characteristics and chloroplast ultrastructure of tea tree[J].Plant Nutrition and Fertilizer Science,2010,16(2):432-438.
[8]申燕,肖家欣,杨慧,等.镁胁迫对“春见”橘橙生长和矿质元素分布及叶片超微结构的影响[J].园艺学报,2011,38(5):849-858.Shen Y,Xiao J X,Yang H,et al.Effects of magnesium stress on growth,distribution of several mineral elements and leaf ultrastructure of‘Harumi’Tangor[J].Acta Horticulturae Sinica,2011,38(5):849-858.
[9]鲍士旦.土壤农化分析(第三版)[M].北京:中国农业出版社,2000.Bao S D.Soil and agricultural chemistry analysis(Third Edition.)[M].Beijing:China Agriculture Press,2000.
[10]王学奎.植物生理生化实验原理和技术(第2版)[M].北京:高等教育出版社,2006:134-136.Wang X K.Principles and techniques of plant physiological and biochemical experiment(2nd Edition.)[M].Beijing:Higher Education Press,2006:134-136.
[11]Garcia-Luis A,Oliveira M E M.Dry matter accumulation in citrus fruit is not limited by transport capacity of the pedicel[J].Annals of Botany,2002,90(6):755-764.
[12]Lianopoulou V,Bosabalidis A M,Patakas A,et al.Effects of chilling stress on leaf morphology,anatomy,ultrastructure,gas exchange,and essential oils in the seasonally dimorphic plant Teucrium polium(Lamiaceae)[J].Acta Physiologiae Plantarum,2014,36(8):2271-2281.
[13]Schaaf G,Schikora A,H?berle J,et al.A putative function for the arabidopsis Fe-phytosiderophore transporter homolog at YSL2 in Fe and Zn homeostasis[J].Plant&Cell Physiology,2005,46(5):762-774.
[14]Moseley J L,Allinger T,Herzog S,et al.Adaptation to Fe-deficiency requires remodeling of the photosynthetic apparatus[J].Embo Journal,2003,21(24):6709-6720.
[15]Oh S A,Park J H,Lee G I,et al.Identification of three genetic loci controlling leaf senescence in Arabidopsis thaliana[J].The Plant Journal,1997,12(3):527-535.
[16]Pestana M,Varennes A D,Abadia J,et al.Differential tolerance to iron deficiency of citrus rootstocks grown in nutrient solution[J].Scientia Horticulturae,2005,104:25-36.
[17]陆志峰,任涛,鲁剑巍,等.缺钾油菜叶片光合速率下降的主导因子及其机理[J].植物营养与肥料学报,2016,22(1):122-131.Lu Z F,Ren T,Lu J W,et al.Main factors and mechanism leading to the decrease of photosynthetic efficiency of oilseed rape exposure to potassium deficiency[J].Journal of Plant Nutrition and Fertilizer,2016,22(1):122-131.
[18]周开兵,郭文武,夏仁学,等.柑橘体细胞杂种砧木对脐橙幼树生长及根和叶中抗氧化酶系活性的影响[J].植物生理学报,2004,40(5):540-544.Zhou K B,Guo W W,Xia R X,et al.Effects of somatic hybrids rootstocks on the growth of young tree and the activities of antioxidant enzymes in leaves and roots of navel orange[J].Plant Physiological Communications,2004,40(5):540-544.
[19]Martínez-Cuenca M R,Iglesias D J,Talón M,et al.Metabolic responses to iron deficiency in roots of Carrizo citrange[Citrus sinensis(L.)Osbeck.×Poncirus trifoliata(L.)Raf][J].Tree Physiology,2013,33(3):320-329.
[20]Pestana M,Varennes A D,Abadía J,et al.Differential tolerance to iron deficiency of citrus rootstocks grown in nutrient solution[J].Scientia Horticulturae,2005,104(1):25-36.
[21]Chen L S,Smith B R,Cheng L.CO2 assimilation,photosynthetic enzymes,and carbohydrates of'concord'grape leaves in response to iron supply[J].Journal of the American Society for Horticultural Science,2004,129(5):738-744.
[22]Arulanantham A R,Rao I M,Terry N.Limiting factors in photosynthesis.VI.Regeneration of ribulose 1,5-bisphosphate limits photosynthesis at low photochemical capacity[J].Plant Physiology,1990,93(4):1466-1475.
[23]王双明,陶诗顺.缺铁胁迫与外源Vc对小白菜生理及品质的影响[J].中国土壤与肥料,2004,(3):26-27.Wang S M,Tao S S.Effects of iron-deficient stress and exogenous Vc on physiological metabolism and quality of vegetables[J].Soil and Fertilizer Sciences in China,2004,(3):26-27.
[24]Li Y,Ren B,Yang X,et al.Chloroplast downsizing under nitrate nutrition restrained mesophyll conductance and photosynthesis in rice(Oryza sativa L.)under drought conditions[J].Plant and Cell Physiology,2012,53(5):892-900.
[25]Weise S E,Carr D J,Bourke A M,et al.The arc,mutants of Arabidopsis,with fewer large chloroplasts have a lower mesophyll conductance[J].Photosynthesis Research,2015,124(1):117-126.
[26]李向东,王晓云,余松烈,等.花生叶片衰老过程中光合性能及细胞微结构变化[J].中国农业科学,2002,35(4):384-389.Li X D,Wang X Y,Yu S L,et al.The changes of photosynthetic properties and cell microstructure in peanut leaves during leaf senescence[J].Scientia Agricultura Sinica,2002,35(4):384-389.
[27]Bernal M,Ramiro M V,Cases R,et al.Excess copper effect on growth,chloroplast ultrastructure,oxygen-evolution activity and chlorophyll fluorescence in Glycine max cell suspensions[J].Physiologia Plantarum,2006,127(2):312-325.
[28]谢深喜,张秋明,熊兴耀,等.水分胁迫对柑橘叶片和根系细胞超微结构的影响[J].湖南农业大学学报(自然科学版),2008,34(2):168-172.Xie S X,Zhang Q M,Xiong X Y,et al.Effect of water stress on leaf and root cell ultra-structure of citrus[J].Journal of Hunan Agricultural University(Natural Sciences),2008,34(2):168-172.
[29]刘磊超,姜存仓,董肖昌,等.硼胁迫对枳橙砧木细根根尖成熟区和幼嫩叶片细胞结构的影响[J].中国农业科学,2015,48(24):4957-4964.Liu L C,Jiang C C,Dong X C,et al.Effects of boron deficiency on cellular structures of maturation zone from root tips and functional leaves from middle and upper plant in trifoliate orange rootstock[J].Scientia Agricultura Sinica,2015,48(24):4957-4964.
[30]肖家欣,齐笑笑,张绍铃.锌和铁缺乏对枳生理指标、矿物质含量及叶片超微结构的影响[J].应用生态学报,2010,21(8):1974-1980.Xiao J X,Qi X X,Zhang S L.Effects of zinc and iron deficiency on physiological indices,mineral contents and leaf ultrastructure of poncirus trifoliate[J].Chinese Journal of Applied Ecology,2010,21(8):1974-1980.
[2]周开兵,郭文武,夏仁学,等.两种柑桔体细胞杂种砧木利用价值和砧穗互作生化机制的探讨[J].园艺学报,2004,31(4):427-432.Zhou K B,Guo W W,Xia R X,et al.Studies on the utilization potential of two somatic hybrid rootstocks and the biochemical mechanism of rootstock-scion interaction[J].Acta Horticulturae Sinica,2004,31(4):427-432.
[3]Sheng O,Song S,Peng S,et al.The effects of low boron on growth,gas exchange,boron concentration and distribution of‘Newhall’navel orange(Citrus sinensis,Osb.)plants grafted on two rootstocks[J].Scientia Horticulturae,2009,121(3):278-283.
[4]Hellín E,Hernández-Cortés J A,Piqueras A,et al.The influence of the iron content on the superoxide dismutase activity and chloroplast ultrastructure of Citrus limon[A].Iron nutrition in soils and plants[M].Springer Netherlands,1995:211-229.
[5]姚允聪,王绍辉,孔云.弱光条件下桃叶片结构及光合特性与叶绿体超微结构变化[J].中国农业科学,2007,40(4):855-863.Yao Y C,Wang S H,Kong Y.Characteristics of photosynthesis mechanism in different peach species under low light intensity[J].Scientia Agricultura Sinica,2007,40(4):855-863.
[6]郭卫东,桑丹,郑建树,等.缺氮对佛手气体交换、叶绿素荧光及叶绿体超微结构的影响[J].浙江大学学报(农业与生命科学版),2009,35(3):307-314.Guo W D,Sang D,Zheng S J,et al.Effects of nitrogen deficiency on the gas exchange,chlorophyll fluorescence and chloroplast ultrastructure in fingered citron[J].Journal of Zhejiang University(Agricultural&Life Sciences),2009,35(3):307-314.
[7]王跃华,张丽霞,孙其远.钙过量对茶树光合特性及叶绿体超微结构的影响[J].植物营养与肥料学报,2010,16(2):432-438.Wang Y H,Zhang L X,Sun Q Y.Effects of excessive calcium fertilization on photosynthetic characteristics and chloroplast ultrastructure of tea tree[J].Plant Nutrition and Fertilizer Science,2010,16(2):432-438.
[8]申燕,肖家欣,杨慧,等.镁胁迫对“春见”橘橙生长和矿质元素分布及叶片超微结构的影响[J].园艺学报,2011,38(5):849-858.Shen Y,Xiao J X,Yang H,et al.Effects of magnesium stress on growth,distribution of several mineral elements and leaf ultrastructure of‘Harumi’Tangor[J].Acta Horticulturae Sinica,2011,38(5):849-858.
[9]鲍士旦.土壤农化分析(第三版)[M].北京:中国农业出版社,2000.Bao S D.Soil and agricultural chemistry analysis(Third Edition.)[M].Beijing:China Agriculture Press,2000.
[10]王学奎.植物生理生化实验原理和技术(第2版)[M].北京:高等教育出版社,2006:134-136.Wang X K.Principles and techniques of plant physiological and biochemical experiment(2nd Edition.)[M].Beijing:Higher Education Press,2006:134-136.
[11]Garcia-Luis A,Oliveira M E M.Dry matter accumulation in citrus fruit is not limited by transport capacity of the pedicel[J].Annals of Botany,2002,90(6):755-764.
[12]Lianopoulou V,Bosabalidis A M,Patakas A,et al.Effects of chilling stress on leaf morphology,anatomy,ultrastructure,gas exchange,and essential oils in the seasonally dimorphic plant Teucrium polium(Lamiaceae)[J].Acta Physiologiae Plantarum,2014,36(8):2271-2281.
[13]Schaaf G,Schikora A,H?berle J,et al.A putative function for the arabidopsis Fe-phytosiderophore transporter homolog at YSL2 in Fe and Zn homeostasis[J].Plant&Cell Physiology,2005,46(5):762-774.
[14]Moseley J L,Allinger T,Herzog S,et al.Adaptation to Fe-deficiency requires remodeling of the photosynthetic apparatus[J].Embo Journal,2003,21(24):6709-6720.
[15]Oh S A,Park J H,Lee G I,et al.Identification of three genetic loci controlling leaf senescence in Arabidopsis thaliana[J].The Plant Journal,1997,12(3):527-535.
[16]Pestana M,Varennes A D,Abadia J,et al.Differential tolerance to iron deficiency of citrus rootstocks grown in nutrient solution[J].Scientia Horticulturae,2005,104:25-36.
[17]陆志峰,任涛,鲁剑巍,等.缺钾油菜叶片光合速率下降的主导因子及其机理[J].植物营养与肥料学报,2016,22(1):122-131.Lu Z F,Ren T,Lu J W,et al.Main factors and mechanism leading to the decrease of photosynthetic efficiency of oilseed rape exposure to potassium deficiency[J].Journal of Plant Nutrition and Fertilizer,2016,22(1):122-131.
[18]周开兵,郭文武,夏仁学,等.柑橘体细胞杂种砧木对脐橙幼树生长及根和叶中抗氧化酶系活性的影响[J].植物生理学报,2004,40(5):540-544.Zhou K B,Guo W W,Xia R X,et al.Effects of somatic hybrids rootstocks on the growth of young tree and the activities of antioxidant enzymes in leaves and roots of navel orange[J].Plant Physiological Communications,2004,40(5):540-544.
[19]Martínez-Cuenca M R,Iglesias D J,Talón M,et al.Metabolic responses to iron deficiency in roots of Carrizo citrange[Citrus sinensis(L.)Osbeck.×Poncirus trifoliata(L.)Raf][J].Tree Physiology,2013,33(3):320-329.
[20]Pestana M,Varennes A D,Abadía J,et al.Differential tolerance to iron deficiency of citrus rootstocks grown in nutrient solution[J].Scientia Horticulturae,2005,104(1):25-36.
[21]Chen L S,Smith B R,Cheng L.CO2 assimilation,photosynthetic enzymes,and carbohydrates of'concord'grape leaves in response to iron supply[J].Journal of the American Society for Horticultural Science,2004,129(5):738-744.
[22]Arulanantham A R,Rao I M,Terry N.Limiting factors in photosynthesis.VI.Regeneration of ribulose 1,5-bisphosphate limits photosynthesis at low photochemical capacity[J].Plant Physiology,1990,93(4):1466-1475.
[23]王双明,陶诗顺.缺铁胁迫与外源Vc对小白菜生理及品质的影响[J].中国土壤与肥料,2004,(3):26-27.Wang S M,Tao S S.Effects of iron-deficient stress and exogenous Vc on physiological metabolism and quality of vegetables[J].Soil and Fertilizer Sciences in China,2004,(3):26-27.
[24]Li Y,Ren B,Yang X,et al.Chloroplast downsizing under nitrate nutrition restrained mesophyll conductance and photosynthesis in rice(Oryza sativa L.)under drought conditions[J].Plant and Cell Physiology,2012,53(5):892-900.
[25]Weise S E,Carr D J,Bourke A M,et al.The arc,mutants of Arabidopsis,with fewer large chloroplasts have a lower mesophyll conductance[J].Photosynthesis Research,2015,124(1):117-126.
[26]李向东,王晓云,余松烈,等.花生叶片衰老过程中光合性能及细胞微结构变化[J].中国农业科学,2002,35(4):384-389.Li X D,Wang X Y,Yu S L,et al.The changes of photosynthetic properties and cell microstructure in peanut leaves during leaf senescence[J].Scientia Agricultura Sinica,2002,35(4):384-389.
[27]Bernal M,Ramiro M V,Cases R,et al.Excess copper effect on growth,chloroplast ultrastructure,oxygen-evolution activity and chlorophyll fluorescence in Glycine max cell suspensions[J].Physiologia Plantarum,2006,127(2):312-325.
[28]谢深喜,张秋明,熊兴耀,等.水分胁迫对柑橘叶片和根系细胞超微结构的影响[J].湖南农业大学学报(自然科学版),2008,34(2):168-172.Xie S X,Zhang Q M,Xiong X Y,et al.Effect of water stress on leaf and root cell ultra-structure of citrus[J].Journal of Hunan Agricultural University(Natural Sciences),2008,34(2):168-172.
[29]刘磊超,姜存仓,董肖昌,等.硼胁迫对枳橙砧木细根根尖成熟区和幼嫩叶片细胞结构的影响[J].中国农业科学,2015,48(24):4957-4964.Liu L C,Jiang C C,Dong X C,et al.Effects of boron deficiency on cellular structures of maturation zone from root tips and functional leaves from middle and upper plant in trifoliate orange rootstock[J].Scientia Agricultura Sinica,2015,48(24):4957-4964.
[30]肖家欣,齐笑笑,张绍铃.锌和铁缺乏对枳生理指标、矿物质含量及叶片超微结构的影响[J].应用生态学报,2010,21(8):1974-1980.Xiao J X,Qi X X,Zhang S L.Effects of zinc and iron deficiency on physiological indices,mineral contents and leaf ultrastructure of poncirus trifoliate[J].Chinese Journal of Applied Ecology,2010,21(8):1974-1980.