核桃树体内矿质营养代谢与SOD酶活性、光合速率的相关性研究
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摘要
本试验以新疆隔年核桃、汾阳绵核桃为试材，通过对其年周期中K、Ca、Mg、Cu、Fe、Mn、Zn七种矿质元素代谢及其与SOD酶活性、光合速率相关性分析，结果显示：
(1)休眠期新疆隔年核桃不同枝类内各矿质元素含量均高于汾阳绵核桃，两品种短果枝中含量又高于其它各类枝。(2)生长期中，枝条和叶片内各元素代谢有不同的变化规律：K含量随着叶龄增大而降低；Ca、Mg含量在枝条内先上升后下降，在叶片中随着叶龄增大而增加；Cu、Fe、Mn、Zn含量在枝条和叶片内变化较稳定。(3)SOD酶活性在两品种幼叶中较高，有利于消除自由基的伤害，老叶中较低，自由基对叶片的破坏能力大；在不同的品种中，新疆隔年核桃SOD酶活性高峰高于汾阳绵核桃，表明不同品种对外界环境影响内在反应不同。(4)核桃两品种的光合速率变化均呈双峰曲线，第二次峰值低于第一次。新疆隔年核桃光合速率高于汾阳绵核桃，这可能是新疆隔年核桃早果、丰产的生理原因之一。(5)在矿质营养、SOD酶活性和光合速率的相关性分析中表明，两品种叶片中SOD酶活性在生长期与Ca、Mg、Cu、Mn、Zn为显著或极显著正相关，与K前期呈负相关，后期为正相关，与Fe前期呈正相关，后期为负相关；矿质元素与光合速率的相关性在品种间存在着一定的差异；新疆隔年核桃结果枝SOD酶活性与光合作用大多呈负相关性，其在发育枝中则6月19日到9月27日为极显著正相关性。(6)不同品种不同枝类和叶片矿质元素间有不同的相关性。在新疆隔年核桃结果枝叶片中Mg与K、Ca、Cu为负相关，其枝条内Cu与Ca、Mg，K与Mn为极显著负相关，Mn与Zn为极显著正相关；在发育枝叶片中Zn与其它元素无显著相关性，其枝条内微量元素间都呈极显著正相关。在汾阳绵核桃结果枝叶片中Zn与其它元素均为不显著负相关，在其枝条内K、Ca、Mg为极显著正相关；在发育枝叶片中，K、Cu与其它元素均为负相关，其枝条内，Ca与Mg、Cu、Fe、Mn、Zn元素间均为负相关。由此可知元素间有一定的拮抗与增效作用。
不同相关性分析表明，供试两品种核桃营养代谢的生理机制除与其本身遗传特性有关外，同时受控于多种生理因素，如酶活性、光合速率及矿质元素种类和含量水平等都是其重要的相关因子。试验结果为品种选优、科学栽培和配方施肥提供了新的理论依据。
The contents of the mineral nutrition(K Ca Mg Cu Fe Mn Zn)in branches and leaves of walnut(including Xinjiang-genian walnut and Fenyang-mian walnut) were measured during the period of dormancy and growth, and the activities of SOD, photosynthetic rate in the leaves were also measured during the period of grow, and analyzed the correlation among them were analyzed . The results were as follows:
(1) During dormant period, the contents of seven mineral nutrition in different branches of Xinjiang-genian walnut were all higher than those of Fenyang-mian walnut .The Contents of mineral nutrition in fertile branch were all higher than those of the others branches.
(2) During the period of growth, the variation of the contents of mineral nutrition in branches and leaves were different regularity, the content of K in leaves and branches dropped with the increasing of the leaves age, those of Ca Mg in the branches increased at former stage and dropped at later stage, whereas in the leaves that increased with the increasing of the leaves age; the variation of the contents of Cu Fe Mn Zn in the branches and leaves were relatively stable.
(3) The activities of SOD in the young leaves were higher than that of the old leaves in varieties walnuts .It could remove the injure of free radicals for the young leaves, and could increase damage for the old leaves. The peak of activities of SOD in Xinjiang-genian walnut was higher than that of Fenyang-mian walnut. It indicated that there were different reactions to the effect of environment in different varieties.
(4) The curves of the change of the photosynthetic rate shown double peak for two varieties, and the first peak was higher than the second peak. The photosynthetic rate of Xinjiang-genian walnut higher than that of Fenyang-mian walnut. Perhaps this is a cause of early fruiting and high yield for Xinjiang-genian walnut.
(5) The correlations among the mineral nutrition, the activities of SOD and the
photosynthetic rate indicate: at the growth period, the positive correlation between the activities of SOD and Ca Mg Cu Mn Zn were significant or every significant in the leaves, and the correlation between the activities of SOD and the content of K wad negative in the early period. And positive in the late time, it is contrary to Fe; In different varieties, the correlation between the contents of mineral nutrition and the photosynthetic rate were different; the correlation between the activity of SOD and photosynthetic rate were negative correlation in fruiting branches of Xinjiang-genian walnut, But that were every positive in extending shoots from 19 June to 29 September.
(6) The correlations among the contents of the mineral nutrition in different braches and leaves of the different varieties were different. The correlation among the contents of Mg and K Ca Cu were negative in the leaves of fertile branches of Xinjiang-genian walnut; The correlation among the contents of Ca Mg and Cu, K and Mn were very negative significant, Mn and Zn was very positively significant in the branches; the correlation between Zn and the other elements were not significant in the leaves of the extending shoots, and the correlation among microelement were all very positive in the branches. The negative correlation among the contents of Zn and the other elements were not significant in the leaves of the fertile branches of Fenyang-mian walnut; the correlations among the contents of K Ca Mg were very significant positive in the branches, and those of the contents of K Cu and Ca Mg Fe Mn Zn were negative in the leaves of extending shoots, and the content of Ca and Mg Cu Fe Mn Zn were all negative in the branches. In other words, there were certain synergism or antagonistic effect among the elements.
The analysis of the different correlation indicate that the physiological mechanisrh of the nutrition metabolize in two varieties tested was relative to genetic characteristic. In addition, it was also controlled many physiological factors. For example, the activities of th
(1)休眠期新疆隔年核桃不同枝类内各矿质元素含量均高于汾阳绵核桃，两品种短果枝中含量又高于其它各类枝。(2)生长期中，枝条和叶片内各元素代谢有不同的变化规律：K含量随着叶龄增大而降低；Ca、Mg含量在枝条内先上升后下降，在叶片中随着叶龄增大而增加；Cu、Fe、Mn、Zn含量在枝条和叶片内变化较稳定。(3)SOD酶活性在两品种幼叶中较高，有利于消除自由基的伤害，老叶中较低，自由基对叶片的破坏能力大；在不同的品种中，新疆隔年核桃SOD酶活性高峰高于汾阳绵核桃，表明不同品种对外界环境影响内在反应不同。(4)核桃两品种的光合速率变化均呈双峰曲线，第二次峰值低于第一次。新疆隔年核桃光合速率高于汾阳绵核桃，这可能是新疆隔年核桃早果、丰产的生理原因之一。(5)在矿质营养、SOD酶活性和光合速率的相关性分析中表明，两品种叶片中SOD酶活性在生长期与Ca、Mg、Cu、Mn、Zn为显著或极显著正相关，与K前期呈负相关，后期为正相关，与Fe前期呈正相关，后期为负相关；矿质元素与光合速率的相关性在品种间存在着一定的差异；新疆隔年核桃结果枝SOD酶活性与光合作用大多呈负相关性，其在发育枝中则6月19日到9月27日为极显著正相关性。(6)不同品种不同枝类和叶片矿质元素间有不同的相关性。在新疆隔年核桃结果枝叶片中Mg与K、Ca、Cu为负相关，其枝条内Cu与Ca、Mg，K与Mn为极显著负相关，Mn与Zn为极显著正相关；在发育枝叶片中Zn与其它元素无显著相关性，其枝条内微量元素间都呈极显著正相关。在汾阳绵核桃结果枝叶片中Zn与其它元素均为不显著负相关，在其枝条内K、Ca、Mg为极显著正相关；在发育枝叶片中，K、Cu与其它元素均为负相关，其枝条内，Ca与Mg、Cu、Fe、Mn、Zn元素间均为负相关。由此可知元素间有一定的拮抗与增效作用。
不同相关性分析表明，供试两品种核桃营养代谢的生理机制除与其本身遗传特性有关外，同时受控于多种生理因素，如酶活性、光合速率及矿质元素种类和含量水平等都是其重要的相关因子。试验结果为品种选优、科学栽培和配方施肥提供了新的理论依据。
The contents of the mineral nutrition(K Ca Mg Cu Fe Mn Zn)in branches and leaves of walnut(including Xinjiang-genian walnut and Fenyang-mian walnut) were measured during the period of dormancy and growth, and the activities of SOD, photosynthetic rate in the leaves were also measured during the period of grow, and analyzed the correlation among them were analyzed . The results were as follows:
(1) During dormant period, the contents of seven mineral nutrition in different branches of Xinjiang-genian walnut were all higher than those of Fenyang-mian walnut .The Contents of mineral nutrition in fertile branch were all higher than those of the others branches.
(2) During the period of growth, the variation of the contents of mineral nutrition in branches and leaves were different regularity, the content of K in leaves and branches dropped with the increasing of the leaves age, those of Ca Mg in the branches increased at former stage and dropped at later stage, whereas in the leaves that increased with the increasing of the leaves age; the variation of the contents of Cu Fe Mn Zn in the branches and leaves were relatively stable.
(3) The activities of SOD in the young leaves were higher than that of the old leaves in varieties walnuts .It could remove the injure of free radicals for the young leaves, and could increase damage for the old leaves. The peak of activities of SOD in Xinjiang-genian walnut was higher than that of Fenyang-mian walnut. It indicated that there were different reactions to the effect of environment in different varieties.
(4) The curves of the change of the photosynthetic rate shown double peak for two varieties, and the first peak was higher than the second peak. The photosynthetic rate of Xinjiang-genian walnut higher than that of Fenyang-mian walnut. Perhaps this is a cause of early fruiting and high yield for Xinjiang-genian walnut.
(5) The correlations among the mineral nutrition, the activities of SOD and the
photosynthetic rate indicate: at the growth period, the positive correlation between the activities of SOD and Ca Mg Cu Mn Zn were significant or every significant in the leaves, and the correlation between the activities of SOD and the content of K wad negative in the early period. And positive in the late time, it is contrary to Fe; In different varieties, the correlation between the contents of mineral nutrition and the photosynthetic rate were different; the correlation between the activity of SOD and photosynthetic rate were negative correlation in fruiting branches of Xinjiang-genian walnut, But that were every positive in extending shoots from 19 June to 29 September.
(6) The correlations among the contents of the mineral nutrition in different braches and leaves of the different varieties were different. The correlation among the contents of Mg and K Ca Cu were negative in the leaves of fertile branches of Xinjiang-genian walnut; The correlation among the contents of Ca Mg and Cu, K and Mn were very negative significant, Mn and Zn was very positively significant in the branches; the correlation between Zn and the other elements were not significant in the leaves of the extending shoots, and the correlation among microelement were all very positive in the branches. The negative correlation among the contents of Zn and the other elements were not significant in the leaves of the fertile branches of Fenyang-mian walnut; the correlations among the contents of K Ca Mg were very significant positive in the branches, and those of the contents of K Cu and Ca Mg Fe Mn Zn were negative in the leaves of extending shoots, and the content of Ca and Mg Cu Fe Mn Zn were all negative in the branches. In other words, there were certain synergism or antagonistic effect among the elements.
The analysis of the different correlation indicate that the physiological mechanisrh of the nutrition metabolize in two varieties tested was relative to genetic characteristic. In addition, it was also controlled many physiological factors. For example, the activities of th
引文
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59．汪洪，褚天铎．植物镁素营养的研究进展．植物学通报，1999，16(3)：245-250
60．应文龙，邵菁，李淑俊．镁离子对巯基修饰类囊体膜H~+-ATP酶的质子传导途径的调节效应．植物生理学报，1991，17(1)：92-98
61．袁清昌，许长成，邹琦．钙以及钙和钙调素拮抗对小麦幼苗LOX活性的影响(简报)．植物生理学通讯，1997，33(4)：266-269
62．赵同科．植物锌营养研究综述与展望．河北农业大学学报，1996，19(1)：102-106
63．何新华．植物中的铁素营养．植物学通报，1992，9(4)：24-28
64．张新生，熊学林，周卫等．苹果钙素营养研究进展．土壤肥料，1999，(4)：3-6
65．蒋德安，饶立华，彭佐权．低K条件下水稻的光合特性．植物生理学报，1988，14(1)：50-55
66．饶立华，蒋德安，薛建明等．K营养对水稻光合器官功能的效应与谷粒产量的影响．植物生理学报，1989，15(2)：191-198
67．李合生主编．植物生理生化实验原理和技术．高等教育出版社．1999
68．刘永梅．低K对番红花光合作用的影响．西南师范大学学报(自然科学版)．1997，22(4)：426-429
69．张往祥，吴家胜，曹福亮．N、P、K三要素对银杏光合性能的影响．江西农业大学学报(自然科学版)．1997，24(6)：810-812
70．王爱国，叶发辉，罗广华．活性氧对花生叶片大分子量DNA的损伤．植物生理学通讯，1993，29(4)：260-262
71．户苅次主编(薛德榕译)．作物的光合作用与物质生产．科学出版社．1979
72．伍泽堂，张刚元．脱落酸、细胞分裂素和丙二醛对SOD活性的影响．植物生理学通讯．1990(4)：30-32
73. Catsky J, Sestak Z. photosynthesis during leaf development. In: Pessarakli M(ed).Handbook of photosynthesis. New York: Marcel Dekker, 1997, 633-660
74. Thomas H,Stoddart J L. Leaf senescence. Ann Rev Plant Physiol,1980,31:83-111
75. Longstreth D J, Nobel P S. Salinity effects on leaf anatomy. Plant Physiol,1979, 63: 700-703
76. Spiller S, Terry N. Limiting factors in photosynthesis. Ⅱ. Iron stress diminishes photochemical capacity by reducing the number of photosynthetic units. Plant Physiol, 1980, 65: 121-125
77. Xu D-Q, Shen Y-K. Light stess: photonhibition of photosynthesis in plants under natural conditions. In: Pessarakli M(d).Handbook of Plant and Crop Stress.New York: Marcel Dekker, 1999,483-497
78. Foyer C H, Lelandais M, Kuner K J. Photo-oxidative stress in plants. Physiol Plant, 1994, 92: 696-717
79. Evans HJ et al. In potassium in biochemistry and physiology. Proc Colloq. Int. potash Inst. 1971,8:13-29
80. Forster H. Capacity of various plant species for the uptake of potassium in pot experiments. Potash Rev. Subject 16.90th suite,1982,9:1-8
81. McCord JM, Fridovich I. Superoxide dismutase: An enzymic function for ekythr ocuprein. J Biol Chem,1960,224:6094-6055
82. Hu As, Guan YL, jiang BF, et al. Study on the mechanism of high temperature damage of citrus and its prevention [J]. South China Fruits, 1998, 27(2): 13-14
83. Gong M, Zhang SX, Li XS et al. The catabolism of lipids and changes of fatty acid composition in the germinating pollen of Pinus yunnanensis and effects of calcium and calmodulin. Chinese J Bot, 1993, 5(2): 159
84. Quinlan J D. Chemical composition of developing and shed fruits of "Laxton's Fortune"apples. J. hort. sci., 1969, 44: 97-106
85. Bangerth E Calcium-related physiological disorders of plants. Annu. Rev. Phytopath al. 1979, 17: 97-122
86. Poovaiah B W. Molecular and cellular aspects of calcium action in plants [J]. Hortscience, 1988,23(2):267-271
87. Leshem YY, Sridhara S, Thompson JE. Involvement of calcium and calmodulin in membrane deterioration during senescence of pea foliage [J].Plant Physiol.,19 84, 75: 329-335
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37．王白坡等．田间条件下砂梨光合作用的研究．园艺学报，1987，14(2)：97-102
38．王文江等．大磨盘柿树光合特性的研究．园艺学报，1983，20(2)：105-110
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48．张绍玲，杨庆山，马香莲等．苹果短枝型品种光合特性研究．果树科学，1991，8(3)：129-134
49．邵邻相，黄伯仲，丁淑静．锌、锰、铁、和铜离子对水稻幼苗生长及SOD酶活性的影响．种子，2001，(6)：16-17
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51．莫开菊，汪兴平．钙与果实采后生理．植物生理学通讯，1994，30(1)：44-47
52．宋纯鹏，梅慧生，储钟稀等．Ca~(2+)对叶绿体中超氧物自由基产生以及由ACC形成乙烯的影响．植物生理学报，1992，18(1)：55-62
53．缪颖，毛洁锜，巢建福等．钙处理对水蜜桃和梨果实中钙含量、细胞膜透性和代谢产物的影响(简报)，植物生理学通讯，1991，27(3)：184-186
54．袁清昌，许长成，邹琦．钙信使系统在百草枯诱导小麦幼苗膜脂过氧化中的作用．植物生理学通讯，1996，32(1)：13-16
55．缪颖，曾广文，朱祝军．钙离子与CaM对火白菜离体叶圆片Ca~(2+)-ATPase和β-1，3-葡聚糖合成酶活性的影响(简报)．植物生理学通讯，1999，35(5)：367
56．龚明，曹宗巽．钙和钙调素对花粉萌发和花粉管生长的调控．植物生理学通讯，1995，31(5)：321-328
57．关军锋，马智宏，张晓敏等．Ca离子与苹果花粉萌发和花粉管生长的关系．果树科学，1999，16(3)：176-179
58．张炜，于振文．麦类作物和水稻的钾素营养生理．山东农业大学学报，1999，25(2)：255—259
59．汪洪，褚天铎．植物镁素营养的研究进展．植物学通报，1999，16(3)：245-250
60．应文龙，邵菁，李淑俊．镁离子对巯基修饰类囊体膜H~+-ATP酶的质子传导途径的调节效应．植物生理学报，1991，17(1)：92-98
61．袁清昌，许长成，邹琦．钙以及钙和钙调素拮抗对小麦幼苗LOX活性的影响(简报)．植物生理学通讯，1997，33(4)：266-269
62．赵同科．植物锌营养研究综述与展望．河北农业大学学报，1996，19(1)：102-106
63．何新华．植物中的铁素营养．植物学通报，1992，9(4)：24-28
64．张新生，熊学林，周卫等．苹果钙素营养研究进展．土壤肥料，1999，(4)：3-6
65．蒋德安，饶立华，彭佐权．低K条件下水稻的光合特性．植物生理学报，1988，14(1)：50-55
66．饶立华，蒋德安，薛建明等．K营养对水稻光合器官功能的效应与谷粒产量的影响．植物生理学报，1989，15(2)：191-198
67．李合生主编．植物生理生化实验原理和技术．高等教育出版社．1999
68．刘永梅．低K对番红花光合作用的影响．西南师范大学学报(自然科学版)．1997，22(4)：426-429
69．张往祥，吴家胜，曹福亮．N、P、K三要素对银杏光合性能的影响．江西农业大学学报(自然科学版)．1997，24(6)：810-812
70．王爱国，叶发辉，罗广华．活性氧对花生叶片大分子量DNA的损伤．植物生理学通讯，1993，29(4)：260-262
71．户苅次主编(薛德榕译)．作物的光合作用与物质生产．科学出版社．1979
72．伍泽堂，张刚元．脱落酸、细胞分裂素和丙二醛对SOD活性的影响．植物生理学通讯．1990(4)：30-32
73. Catsky J, Sestak Z. photosynthesis during leaf development. In: Pessarakli M(ed).Handbook of photosynthesis. New York: Marcel Dekker, 1997, 633-660
74. Thomas H,Stoddart J L. Leaf senescence. Ann Rev Plant Physiol,1980,31:83-111
75. Longstreth D J, Nobel P S. Salinity effects on leaf anatomy. Plant Physiol,1979, 63: 700-703
76. Spiller S, Terry N. Limiting factors in photosynthesis. Ⅱ. Iron stress diminishes photochemical capacity by reducing the number of photosynthetic units. Plant Physiol, 1980, 65: 121-125
77. Xu D-Q, Shen Y-K. Light stess: photonhibition of photosynthesis in plants under natural conditions. In: Pessarakli M(d).Handbook of Plant and Crop Stress.New York: Marcel Dekker, 1999,483-497
78. Foyer C H, Lelandais M, Kuner K J. Photo-oxidative stress in plants. Physiol Plant, 1994, 92: 696-717
79. Evans HJ et al. In potassium in biochemistry and physiology. Proc Colloq. Int. potash Inst. 1971,8:13-29
80. Forster H. Capacity of various plant species for the uptake of potassium in pot experiments. Potash Rev. Subject 16.90th suite,1982,9:1-8
81. McCord JM, Fridovich I. Superoxide dismutase: An enzymic function for ekythr ocuprein. J Biol Chem,1960,224:6094-6055
82. Hu As, Guan YL, jiang BF, et al. Study on the mechanism of high temperature damage of citrus and its prevention [J]. South China Fruits, 1998, 27(2): 13-14
83. Gong M, Zhang SX, Li XS et al. The catabolism of lipids and changes of fatty acid composition in the germinating pollen of Pinus yunnanensis and effects of calcium and calmodulin. Chinese J Bot, 1993, 5(2): 159
84. Quinlan J D. Chemical composition of developing and shed fruits of "Laxton's Fortune"apples. J. hort. sci., 1969, 44: 97-106
85. Bangerth E Calcium-related physiological disorders of plants. Annu. Rev. Phytopath al. 1979, 17: 97-122
86. Poovaiah B W. Molecular and cellular aspects of calcium action in plants [J]. Hortscience, 1988,23(2):267-271
87. Leshem YY, Sridhara S, Thompson JE. Involvement of calcium and calmodulin in membrane deterioration during senescence of pea foliage [J].Plant Physiol.,19 84, 75: 329-335