缺钙大白菜的光合特征
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摘要
为系统研究缺钙对大白菜幼苗生长的影响,本文采用水培法分析了不同缺钙程度下,大白菜幼苗的生长、生化和光合指标的特征性变化。研究结果显示,与对照Hoagland完全营养液的100%Ca(4 mmol·L~(-1) Ca~(2+))相比,50%Ca(2 mmol·L~(-1) Ca~(2+))培养4周虽然没有显著抑制大白菜幼苗的生长和净光合速率(P_n),但其叶绿素含量已显著下降;随着缺钙程度的加重,大白菜幼苗的整株鲜重和P_n逐渐下降,叶片叶绿素a/b比值逐渐增大,叶绿素含量和可溶性糖含量则呈先降低后增大的趋势。缺钙大白菜的叶绿素荧光动力学ΔV_t和ΔW_t曲线上最明显的特征是在100 ms处明显出现一个凹峰(G相);此外在缺钙大白菜ΔW_t曲线的K相上(300μs)也出现了特征性的凹峰,且随着缺钙程度的加重,凹峰值下降。两处凹峰的出现说明缺钙导致供体侧电子供应相对减少,但光反应中心的结构和功能未受影响。缺钙条件下,大白菜叶片叶绿素荧光参数的主要特征是由于放氧复合体电子供应能力减弱,导致到达最大荧光的时间(T_(fm))显著延长,单个反应中心的光化学效率(ABS/RC、TR_o/RC、ET_o/RC)下降,质体醌库(S_m)相对增大。上述结果为了解缺钙对大白菜幼苗的胁迫效应和伤害机制以及大白菜缺钙的诊断提供了数据支撑。
To systematically investigate the effects of calcium deficiency on the growth of Chinese cabbage,the characteristic changes of growth,biochemical,and photosynthetic indexes of Chinese cabbage seedlings under calcium deficiency were analyzed using a solution culture method.The results showed that 50%Ca(2 mmol·L~(-1)Ca~(2+))had no significant inhibition on the growth and net photosynthetic rate(P_n)during a 4-week treatment compared with the control(100%Ca:4 mmol·L~(-1) Ca~(2+)in Hoagland’s nutrient solution),while its chlorophyll content was significantly lower than that of the control.With the aggravation of calcium deficiency,the fresh weight of the whole plant and P_n decreased gradually,and the ratio of chlorophyll a/b increased gradually.However,the chlorophyll content and soluble sugar content decreased at first and then increased.The most obvious feature on the chlorophyll fluorescenceΔV_t andΔW_t curve of calcium-deficient cabbage was a clear concave peak at 100 ms(G-phase).In addition,a characteristic concave peak appeared at the K phase of theΔW_(t )curve of the calcium-deficient seedlings,and the concave peak value decreased with the aggravation of calcium deficiency.The two concave peaks at the K phase and G-phase indicated that calcium deficiency led to a relative decrease in electron supply on the donor side,but the structure and function of the reaction center were not affected.Under calcium deficient condition,the main feature of the chlorophyll fluorescence parameters was the weakening of the electron-supply capacity of the oxygen-evolving complex,which led to a significant extension of the time to reach maximal fluorescence intensity F_m(T_(fm)),a decreased photochemical efficiency of a single reaction center(ABS/RC,TR_o/RC,ET_o/RC),and a relatively larger plastoquinone pool.Taken together,this study contributes to our understanding of the effects of calcium deficiency on Chinese cabbage,and provides valuable information for the diagnosis of calcium deficiency of Chinese cabbage.
To systematically investigate the effects of calcium deficiency on the growth of Chinese cabbage,the characteristic changes of growth,biochemical,and photosynthetic indexes of Chinese cabbage seedlings under calcium deficiency were analyzed using a solution culture method.The results showed that 50%Ca(2 mmol·L~(-1)Ca~(2+))had no significant inhibition on the growth and net photosynthetic rate(P_n)during a 4-week treatment compared with the control(100%Ca:4 mmol·L~(-1) Ca~(2+)in Hoagland’s nutrient solution),while its chlorophyll content was significantly lower than that of the control.With the aggravation of calcium deficiency,the fresh weight of the whole plant and P_n decreased gradually,and the ratio of chlorophyll a/b increased gradually.However,the chlorophyll content and soluble sugar content decreased at first and then increased.The most obvious feature on the chlorophyll fluorescenceΔV_t andΔW_t curve of calcium-deficient cabbage was a clear concave peak at 100 ms(G-phase).In addition,a characteristic concave peak appeared at the K phase of theΔW_(t )curve of the calcium-deficient seedlings,and the concave peak value decreased with the aggravation of calcium deficiency.The two concave peaks at the K phase and G-phase indicated that calcium deficiency led to a relative decrease in electron supply on the donor side,but the structure and function of the reaction center were not affected.Under calcium deficient condition,the main feature of the chlorophyll fluorescence parameters was the weakening of the electron-supply capacity of the oxygen-evolving complex,which led to a significant extension of the time to reach maximal fluorescence intensity F_m(T_(fm)),a decreased photochemical efficiency of a single reaction center(ABS/RC,TR_o/RC,ET_o/RC),and a relatively larger plastoquinone pool.Taken together,this study contributes to our understanding of the effects of calcium deficiency on Chinese cabbage,and provides valuable information for the diagnosis of calcium deficiency of Chinese cabbage.
引文
Adam MS,Issa AA(2000).Effect of manganese and calcium deficiency on the growth and oxygen exchange of Scenedesmus intermedius cultured for successive generations.Folia Microbiol,45(4):353-358
Chatzistathis TA,Papadakis IE,Therios IN,et al(2011).Is chlorophyll fluorescence technique a useful tool to assess manganese deficiency and toxicity stress in olive plants?J Plant Nutr,34:98-114
Chen C,Kazimir J,Cheniae GM(1995).Calcium modulates the photoassembly of photosystem II(Mn)4-clusters by preventing ligation of nonfunctional high-valency states of manganese.Biochemistry,34(41):13511-13526
Clarkson DT,Hanson JB(1980).The mineral nutrition of higher plants.Annu Rev Plant Physiol,31:239-298
Conway WS,Sams CE,Kelman A(1994).Enhancing the natural resistance of plant tissues to postharvest diseases through calcium applications.Hort Sci,21(1):66-69
Felix KCDS,Silva CLD,Oliveira WJD,et al(2017).Calcium-mediated reduction of soft rot disease in Chinese cabbage.Eur J Plant Pathol,147:73-84
Hepler PK,Wayne RO(1985).Calcium and plant development.Annu Rev Plant Physiol,36:397-439
Huang H,Chen L,Liu XQ,et al(2008).Absorption and transfer of light and photoreduction activities of spinach chloroplasts under calcium deficiency:promotion by cerium.Biol Trace Elem Res,122(2):157-167
Kalaji HM,Oukarroum A,Alexandrov V,et al(2014).Identification of nutrient deficiency in maize and tomato plants by in vivo chlorophyll a fluorescence measurements.Plant Physiol Biochem,81:16-25
Li H,Liu SQ,Wang Y,et al(2013).Effects of calcium on photosynthetic characteristics and quality of garlic.Acta Hortic Sin,40(6):1169-1177(in Chinese with English abstract)[李贺,刘世琦,王越等(2013).钙对水培大蒜光合特性和品质的影响.园艺学报,40(6):1169-1177]
Li Z,Tan XF,Lu K,et al(2017).The effect of Ca Cl2 on calcium content,photosynthesis,and chlorophyll fluorescence of tung tree seedlings under drought conditions.Photosynthetica,55(3):553-560
Liang WJ,Wang ML,Ai XZ(2009).The role of calcium in regulating photosynthesis and related physiological indexes of cucumber seedlings under low light intensity and suboptimal temperature stress.Sci Hortic,123(1):34-38
Liao RY,Yin LX,Jin G,et al(2008).Effect of calcium concentration on chloroplast cytochrome content and ultrastructure of loquat seedling leaves.Fujian J Agric Sci,23(3):302-305(in Chinese with English abstract)[廖汝玉,尹兰香,金光等(2008).不同浓度钙处理对枇杷小苗叶片细胞超微结构及叶绿体色素含量的影响.福建农业学报,23(3):302-305]
Liu C,Pan BF,Cao WQ,et al(2008).Influences of calcium deficiency and cerium on growth of spinach plants.Biol Trace Elem Res,121(3):266-275
Liu HC,Liu WP,Song SW,et al(2012).Effect of calcium nutrient on calcium distribution and ultrastructure of cell and chloroplast in bunching onion leaf.Appl Mech Mater,142(3):111-115
Liu YF,Han XR,Zhan XM,et al(2013).Regulation of calcium on peanut photosynthesis under low night temperature stress.J Integr Agr,12(12):2172-2178
Miao Y,Wu BH,Zeng GW,et al(2000).Relation between changes in ultrastructure and composition of cell wall and calcium-related physiological disorder of Chinese cabbage(Brassica campestris ssp.pekinensis).Acta Phytophysiol Sin,26(2):111-116(in Chinese with English abstract)[缪颖,伍炳华,曾广文等(2000).缺钙诱发大白菜干烧心与细胞壁结构组分变化的关系.植物生理学报,26(2):111-116]
Qiu NW,Wang XS,Yang FB,et al(2016).Fast extraction and precise determination of chlorophyll.Chin Bull Bot,51(5):667-678(in Chinese with English abstract)[邱念伟,王修顺,杨发斌等(2016).叶绿素的快速提取与精密测定.植物学报,51(5):667-678]
Ridolfi M,Roupsard O,Garrec JP,et al(1996).Effects of a calcium deficiency on stomatal conductance and photosynthetic activity of Quercus robur seedlings grown on nutrient solution.Ann Sci For,53:325-335
Strasser RJ,Srivastava A,Tsimilli-Michael M(2000).The fluorescence transient as a tool to characterise and screen photosynthetic samples.In:Probing Photosynthesis:Mechanisms,Regulation and Adaptation.Yunus M,Pathre U,Mohanty E(eds).London:Taylor&Francis,445-483
Tan W,Meng QW,Brestic M,et al(2011).Photosynthesis is improved by exogenous calcium in heat-stressed tobacco plants.J Plant Physiol,168(17):2063-2071
Veierskov B,MeravyL(1985).Changes in carbohydrate composition in wheat and pea seedlings induced by calcium deficiency.Plant Physiol,79(1):315-317
Wang SF,Zhang Y,Shen ZY(1996).Morphological,structural,physiological and biochemical changes during the develoment of tipburn symptoms in Chinese cabbage.Acta Hortic Sin,23(1):37-44(in Chinese with English abstract)[王淑芬,张仪,沈征言(1996).大白菜干烧心病的形态结构及生理生化变化.园艺学报,23(1):37-44]
Yang ZL,Li LB,Xu YN,et al(2000).Effect of phosphatidyl glycerol(PG)on oxygen release activity in calcium-deficient PSII.Chinese Sci Bull,45(9):950-955(in Chinese)[阳振乐,李良璧,许亦农等(2000).磷脂酰甘油对缺钙光系统II放氧活性的影响.科学通报,45(9):950-955]
Zhang GX,Liu YF,Ni Y,et al(2014).Exogenous calcium alleviates low night temperature stress on the photosynthetic apparatus of tomato leaves.PLo S One,9(5):e97322
Zhao SE,Xing JM,Li DZ(1982).The relationship between head tipburn and calcium deficiency of Chinese cabbage(Brassica pekinensis Rupr.).Acta Hortic Sin,9(1):33-40(in Chinese with English abstract)[赵素娥,邢金铭,李得众(1982).大白菜“干心”病的发生与缺钙的关系.园艺学报,9(1):33-40]
Zhou W,Lin B(1995).Research progress in calcium nutrition mechanism of plant.Prog Soil Sci,23(2):12-17(in Chinese)[周卫,林葆(1995).植物钙素营养机理研究进展.土壤学进展,23(2):12-17]
Zheng Y,Chen ZJ(2015).Organellar calcium signaling in plants.Plant Physiol J,51(8):1195-1203(in Chinese with English abstract)[郑远,陈兆进(2015).植物细胞器钙信号研究进展.植物生理学报,51(8):1195-1203]
Chatzistathis TA,Papadakis IE,Therios IN,et al(2011).Is chlorophyll fluorescence technique a useful tool to assess manganese deficiency and toxicity stress in olive plants?J Plant Nutr,34:98-114
Chen C,Kazimir J,Cheniae GM(1995).Calcium modulates the photoassembly of photosystem II(Mn)4-clusters by preventing ligation of nonfunctional high-valency states of manganese.Biochemistry,34(41):13511-13526
Clarkson DT,Hanson JB(1980).The mineral nutrition of higher plants.Annu Rev Plant Physiol,31:239-298
Conway WS,Sams CE,Kelman A(1994).Enhancing the natural resistance of plant tissues to postharvest diseases through calcium applications.Hort Sci,21(1):66-69
Felix KCDS,Silva CLD,Oliveira WJD,et al(2017).Calcium-mediated reduction of soft rot disease in Chinese cabbage.Eur J Plant Pathol,147:73-84
Hepler PK,Wayne RO(1985).Calcium and plant development.Annu Rev Plant Physiol,36:397-439
Huang H,Chen L,Liu XQ,et al(2008).Absorption and transfer of light and photoreduction activities of spinach chloroplasts under calcium deficiency:promotion by cerium.Biol Trace Elem Res,122(2):157-167
Kalaji HM,Oukarroum A,Alexandrov V,et al(2014).Identification of nutrient deficiency in maize and tomato plants by in vivo chlorophyll a fluorescence measurements.Plant Physiol Biochem,81:16-25
Li H,Liu SQ,Wang Y,et al(2013).Effects of calcium on photosynthetic characteristics and quality of garlic.Acta Hortic Sin,40(6):1169-1177(in Chinese with English abstract)[李贺,刘世琦,王越等(2013).钙对水培大蒜光合特性和品质的影响.园艺学报,40(6):1169-1177]
Li Z,Tan XF,Lu K,et al(2017).The effect of Ca Cl2 on calcium content,photosynthesis,and chlorophyll fluorescence of tung tree seedlings under drought conditions.Photosynthetica,55(3):553-560
Liang WJ,Wang ML,Ai XZ(2009).The role of calcium in regulating photosynthesis and related physiological indexes of cucumber seedlings under low light intensity and suboptimal temperature stress.Sci Hortic,123(1):34-38
Liao RY,Yin LX,Jin G,et al(2008).Effect of calcium concentration on chloroplast cytochrome content and ultrastructure of loquat seedling leaves.Fujian J Agric Sci,23(3):302-305(in Chinese with English abstract)[廖汝玉,尹兰香,金光等(2008).不同浓度钙处理对枇杷小苗叶片细胞超微结构及叶绿体色素含量的影响.福建农业学报,23(3):302-305]
Liu C,Pan BF,Cao WQ,et al(2008).Influences of calcium deficiency and cerium on growth of spinach plants.Biol Trace Elem Res,121(3):266-275
Liu HC,Liu WP,Song SW,et al(2012).Effect of calcium nutrient on calcium distribution and ultrastructure of cell and chloroplast in bunching onion leaf.Appl Mech Mater,142(3):111-115
Liu YF,Han XR,Zhan XM,et al(2013).Regulation of calcium on peanut photosynthesis under low night temperature stress.J Integr Agr,12(12):2172-2178
Miao Y,Wu BH,Zeng GW,et al(2000).Relation between changes in ultrastructure and composition of cell wall and calcium-related physiological disorder of Chinese cabbage(Brassica campestris ssp.pekinensis).Acta Phytophysiol Sin,26(2):111-116(in Chinese with English abstract)[缪颖,伍炳华,曾广文等(2000).缺钙诱发大白菜干烧心与细胞壁结构组分变化的关系.植物生理学报,26(2):111-116]
Qiu NW,Wang XS,Yang FB,et al(2016).Fast extraction and precise determination of chlorophyll.Chin Bull Bot,51(5):667-678(in Chinese with English abstract)[邱念伟,王修顺,杨发斌等(2016).叶绿素的快速提取与精密测定.植物学报,51(5):667-678]
Ridolfi M,Roupsard O,Garrec JP,et al(1996).Effects of a calcium deficiency on stomatal conductance and photosynthetic activity of Quercus robur seedlings grown on nutrient solution.Ann Sci For,53:325-335
Strasser RJ,Srivastava A,Tsimilli-Michael M(2000).The fluorescence transient as a tool to characterise and screen photosynthetic samples.In:Probing Photosynthesis:Mechanisms,Regulation and Adaptation.Yunus M,Pathre U,Mohanty E(eds).London:Taylor&Francis,445-483
Tan W,Meng QW,Brestic M,et al(2011).Photosynthesis is improved by exogenous calcium in heat-stressed tobacco plants.J Plant Physiol,168(17):2063-2071
Veierskov B,MeravyL(1985).Changes in carbohydrate composition in wheat and pea seedlings induced by calcium deficiency.Plant Physiol,79(1):315-317
Wang SF,Zhang Y,Shen ZY(1996).Morphological,structural,physiological and biochemical changes during the develoment of tipburn symptoms in Chinese cabbage.Acta Hortic Sin,23(1):37-44(in Chinese with English abstract)[王淑芬,张仪,沈征言(1996).大白菜干烧心病的形态结构及生理生化变化.园艺学报,23(1):37-44]
Yang ZL,Li LB,Xu YN,et al(2000).Effect of phosphatidyl glycerol(PG)on oxygen release activity in calcium-deficient PSII.Chinese Sci Bull,45(9):950-955(in Chinese)[阳振乐,李良璧,许亦农等(2000).磷脂酰甘油对缺钙光系统II放氧活性的影响.科学通报,45(9):950-955]
Zhang GX,Liu YF,Ni Y,et al(2014).Exogenous calcium alleviates low night temperature stress on the photosynthetic apparatus of tomato leaves.PLo S One,9(5):e97322
Zhao SE,Xing JM,Li DZ(1982).The relationship between head tipburn and calcium deficiency of Chinese cabbage(Brassica pekinensis Rupr.).Acta Hortic Sin,9(1):33-40(in Chinese with English abstract)[赵素娥,邢金铭,李得众(1982).大白菜“干心”病的发生与缺钙的关系.园艺学报,9(1):33-40]
Zhou W,Lin B(1995).Research progress in calcium nutrition mechanism of plant.Prog Soil Sci,23(2):12-17(in Chinese)[周卫,林葆(1995).植物钙素营养机理研究进展.土壤学进展,23(2):12-17]
Zheng Y,Chen ZJ(2015).Organellar calcium signaling in plants.Plant Physiol J,51(8):1195-1203(in Chinese with English abstract)[郑远,陈兆进(2015).植物细胞器钙信号研究进展.植物生理学报,51(8):1195-1203]