钙对根际淹水胁迫下辣椒幼苗生长及生理代谢的影响
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摘要
自然条件下某些排水不良或地下水位过高的土壤和低洼、沼泽地带，发生洪水或暴雨之后，常会出现因水分过多而对植物造成危害，影响植株正常生长发育，成为困扰农业生产的限制性因子。提高植株在水淹胁迫下的耐性和抗性对农业生产具有重要意义。Ca~(2+)作为植物生长所必需的大量元素和偶联胞外信号与胞内生理反应的第二信使，在逆境胁迫下细胞内含量显著提高，能够将外界环境胁迫信号转导到细胞内，与胞内CaM形成Ca~(2+)·CaM复合物，通过调节基因表达诱导一些生理和生化代谢途径的改变，可缓解逆境胁迫对植物的伤害。所以，研究钙对根际淹水胁迫下辣椒幼苗生长及生理代谢的影响具有重要的理论价值和实践意义。
本文以贵州省地方品种花溪党武辣椒(Capsicum frutescent L．)为试材，通过浸种和叶面喷施不同浓度Ca~(2+)处理，研究根际淹水胁迫下Ca~(2+)对辣椒幼苗生长的影响，探索最佳Ca~(2+)处理方法，以期最大程度上减轻水淹胁迫对辣椒植株的伤害。然后以筛选出的最佳Ca~(2+)处理方法(10mmol·L~(-1)Ca~(2+)浸种)，来研究Ca~(2+)对水淹胁迫下辣椒幼苗根系呼吸代谢，叶片活性氧代谢，根、茎、叶碳水化合物代谢的影响及其与水淹耐性的关系，探讨Ca~(2+)在辣椒植株适应水淹胁迫过程中的生理调节功能。主要研究结果如下：
钙素对水淹胁迫下辣椒幼苗的调控效应具有两面性，一定浓度范围内成正效应，过高反而成负效应。本试验结果表明：Ca~(2+)减轻水淹对辣椒幼苗伤害的最佳处理方法为：10mmol·L~(-1) Ca~(2+)浸种处理。
耐涝性指标测定结果表明外源Ca~(2+)处理能有效抑制水淹胁迫对叶绿素的破坏，膜透性的升高和根系活力的减弱，相对提高了幼苗地上部、地下部干鲜重，减轻水淹胁迫下根系的损伤，表明了外源Ca~(2+)处理能提高辣椒幼苗的耐涝性。
抗氧化酶活性及非酶抗氧化剂含量测定显示Ca~(2+)处理后的辣椒幼苗叶片具有维持较高水平活性氧代谢作用，有利于诱导SOD等抗氧化酶在细胞内的生物合成以及在水淹处理过程中有助于保持其酶活性的作用，对ASA和GSH等非酶抗氧化剂也有明显的保护作用。从而提高植物对自由基的清除能力，减轻了膜系统的过氧化损伤，这可能是Ca~(2+)提高辣椒幼苗耐涝性生理生化基础之一。
外源Ca~(2+)对水淹胁迫下辣椒幼苗的伤害有显著的缓解效应，可改善根系的生长状况，显著地提高水淹胁迫下辣椒幼苗的鲜重和干重；与单纯水淹胁迫相比，外源Ca~(2+)处理能够提高ADH活性，降低LDH活性，避免乳酸和乙醛的积累，同时使植株保持较高的MDH和SDH活性，进行一定的有氧呼吸代谢。表明外源Ca~(2+)可通过调节辣椒幼苗根系内呼吸代谢来缓解水淹胁迫对植株的伤害。
外源Ca~(2+)处理能够提高辣椒幼苗根系、茎部和叶片可溶性糖、蔗糖和果糖含量，降低根系淀粉和葡萄糖的含量，能够改善辣椒植株地上部和地下部碳水化合物的比例，进而促进辣椒幼苗形成新的碳水化合物代谢，来适应水淹逆境。表明在水淹胁迫下，Ca~(2+)参与调节辣椒幼苗碳水化合物的含量，对提高幼苗对水淹胁迫的适应性起着重要作用。
Under natural condition, after having the flood or the rainstorm, certain low-lying soil and bog region of imperfectly drained or high subsoil water level, because of the excessive water contents are often leading to harm to the plant, and affecting the normal growth of plants, and becoming the restrictive factor. So it is very significant to enhance plant's patience and the resistance under flooding stress to the agricultural production. As a major element necessary for plant growth and a second messenger of coupling extra-cellular signal and intracellular physiological reactions, calcium (Ca~(2+)) plays a crucial role in regulating physiological reactions under environment stress. It has been known that [Ca~(2+)]_(cyt) increase rapidly under environmental stress and Ca~(2+)·CaM compound formation after Ca~(2+) combined with target enzyme CaM. It could relieve the harm of stress to the crops by regulating gene expression and inducing change of physiological and biochemical metabolism under environmental stress. Plants have evolved mechanisms for flood stress and adjusting their cellular metabolism to adapt for environmental stress. Therefore, it has the important theory value and the practice significance to research the effect of calcium on growth and physiological metabolism of pepper seedlings under root-zone flood stress.
The research used HuaxiDangwu hot pepper of Guizhou Province as the material, through soaking seeds and spurting the leaf of the different consistence Ca~(2+), researching the effect of Ca~(2+) on growth of pepper seedlings under root-zone flood stress, exploration the best Ca~(2+) processing method, reduced the injury of pepper seedling under flooding stress . Then useing the best processing method of Ca~(2+) (10mmol·L~(-1) Ca~(2+) soaking seeds), studing the influence about Ca~(2+) to the hot pepper seedling root system respiratory metabolism, leaf ROS metabolism, the carbohydrate metabolism of roots, stems, leaves and the relation under the flood stress, discussing the physiological regulation function about Ca~(2+) in the adaptation flooding stress process of pepper plant. Main research results were as follows:
The regulative effect of calcium element to pepper seedling under flood stress has tow sides. On the one hand it is advantage in the certain consistence scope; on the other hand it is negative when the consistence is excessively high. This test result indicated: the best processing method of Ca~(2+) reduces the injury of flood to the hot pepper seedling is: 10mmol·L~(-1) Ca~(2+) soaking seeds processing.
The measuration's result indicated that the extraneous Ca~(2+) processing can suppress effectively to the chlorophyll destruction, membrane penetrable elevation and root activity being weaken under flood stress, and relatively enhanced the fresh weight and dry weight of seedling, reduced root system's damage. These indicated the extraneous Ca~(2+) processing could enhance the resistance of hot pepper seedling to overhead flooding injury.
The measuration about oxidation resistance enzyme activity and the non-enzyme oxidation inhibitor content demonstrate that after the Ca~(2+) processing, the hot pepper seedling leaf has the function maintain high level ROS metabolism, its mechanism possibly is advantageous to induce biosynthesis and keep activity about anti-oxidase enzyme of SOD et al., and non-enzyme oxidation inhibitor of ASA and GSH. Thus enhances the ability of plant to eliminate the free radical, reduced the membrane system's peroxidation damage. This possibly is a physiology biochemistry foundation of Ca~(2+) could enhance the resistance of hot pepper seedling to overhead flooding injury.
Exogenous Ca~(2+) application in nutrient solution could relieve the harm caused by flood stress and result in improved growth of roots, increased fresh and dry weight of pepper seedlings. Compared with the seedlings with flood alone treatment, the activities of SDH, MDH and ADH were promoted, while LDH activity was inhibited, accumulations of lactate and acetaldehyde in pepper seedling roots under exogenous application Ca~(2+) treatment. It indicated that Ca~(2+) could relieve the harm of flood stress, through regulating respiratory metabolism of roots.
Exogenous Ca~(2+) application treatment could increase the contents of and solubility sugar, cane sugar and fruit sugar in roots, stems and leaves of pepper seedlings, decreased the content of starch and glucose in pepper seedling roots, which could improve in the ratio of root carbohydrate concentration to leaves carbohydrate concentration, and promote the appearance of carbohydrate metabolism; adapting the flood stress. It indicated that Ca~(2+) could adjust carbohydrate contents and relieve the harm of flood stress.
本文以贵州省地方品种花溪党武辣椒(Capsicum frutescent L．)为试材，通过浸种和叶面喷施不同浓度Ca~(2+)处理，研究根际淹水胁迫下Ca~(2+)对辣椒幼苗生长的影响，探索最佳Ca~(2+)处理方法，以期最大程度上减轻水淹胁迫对辣椒植株的伤害。然后以筛选出的最佳Ca~(2+)处理方法(10mmol·L~(-1)Ca~(2+)浸种)，来研究Ca~(2+)对水淹胁迫下辣椒幼苗根系呼吸代谢，叶片活性氧代谢，根、茎、叶碳水化合物代谢的影响及其与水淹耐性的关系，探讨Ca~(2+)在辣椒植株适应水淹胁迫过程中的生理调节功能。主要研究结果如下：
钙素对水淹胁迫下辣椒幼苗的调控效应具有两面性，一定浓度范围内成正效应，过高反而成负效应。本试验结果表明：Ca~(2+)减轻水淹对辣椒幼苗伤害的最佳处理方法为：10mmol·L~(-1) Ca~(2+)浸种处理。
耐涝性指标测定结果表明外源Ca~(2+)处理能有效抑制水淹胁迫对叶绿素的破坏，膜透性的升高和根系活力的减弱，相对提高了幼苗地上部、地下部干鲜重，减轻水淹胁迫下根系的损伤，表明了外源Ca~(2+)处理能提高辣椒幼苗的耐涝性。
抗氧化酶活性及非酶抗氧化剂含量测定显示Ca~(2+)处理后的辣椒幼苗叶片具有维持较高水平活性氧代谢作用，有利于诱导SOD等抗氧化酶在细胞内的生物合成以及在水淹处理过程中有助于保持其酶活性的作用，对ASA和GSH等非酶抗氧化剂也有明显的保护作用。从而提高植物对自由基的清除能力，减轻了膜系统的过氧化损伤，这可能是Ca~(2+)提高辣椒幼苗耐涝性生理生化基础之一。
外源Ca~(2+)对水淹胁迫下辣椒幼苗的伤害有显著的缓解效应，可改善根系的生长状况，显著地提高水淹胁迫下辣椒幼苗的鲜重和干重；与单纯水淹胁迫相比，外源Ca~(2+)处理能够提高ADH活性，降低LDH活性，避免乳酸和乙醛的积累，同时使植株保持较高的MDH和SDH活性，进行一定的有氧呼吸代谢。表明外源Ca~(2+)可通过调节辣椒幼苗根系内呼吸代谢来缓解水淹胁迫对植株的伤害。
外源Ca~(2+)处理能够提高辣椒幼苗根系、茎部和叶片可溶性糖、蔗糖和果糖含量，降低根系淀粉和葡萄糖的含量，能够改善辣椒植株地上部和地下部碳水化合物的比例，进而促进辣椒幼苗形成新的碳水化合物代谢，来适应水淹逆境。表明在水淹胁迫下，Ca~(2+)参与调节辣椒幼苗碳水化合物的含量，对提高幼苗对水淹胁迫的适应性起着重要作用。
Under natural condition, after having the flood or the rainstorm, certain low-lying soil and bog region of imperfectly drained or high subsoil water level, because of the excessive water contents are often leading to harm to the plant, and affecting the normal growth of plants, and becoming the restrictive factor. So it is very significant to enhance plant's patience and the resistance under flooding stress to the agricultural production. As a major element necessary for plant growth and a second messenger of coupling extra-cellular signal and intracellular physiological reactions, calcium (Ca~(2+)) plays a crucial role in regulating physiological reactions under environment stress. It has been known that [Ca~(2+)]_(cyt) increase rapidly under environmental stress and Ca~(2+)·CaM compound formation after Ca~(2+) combined with target enzyme CaM. It could relieve the harm of stress to the crops by regulating gene expression and inducing change of physiological and biochemical metabolism under environmental stress. Plants have evolved mechanisms for flood stress and adjusting their cellular metabolism to adapt for environmental stress. Therefore, it has the important theory value and the practice significance to research the effect of calcium on growth and physiological metabolism of pepper seedlings under root-zone flood stress.
The research used HuaxiDangwu hot pepper of Guizhou Province as the material, through soaking seeds and spurting the leaf of the different consistence Ca~(2+), researching the effect of Ca~(2+) on growth of pepper seedlings under root-zone flood stress, exploration the best Ca~(2+) processing method, reduced the injury of pepper seedling under flooding stress . Then useing the best processing method of Ca~(2+) (10mmol·L~(-1) Ca~(2+) soaking seeds), studing the influence about Ca~(2+) to the hot pepper seedling root system respiratory metabolism, leaf ROS metabolism, the carbohydrate metabolism of roots, stems, leaves and the relation under the flood stress, discussing the physiological regulation function about Ca~(2+) in the adaptation flooding stress process of pepper plant. Main research results were as follows:
The regulative effect of calcium element to pepper seedling under flood stress has tow sides. On the one hand it is advantage in the certain consistence scope; on the other hand it is negative when the consistence is excessively high. This test result indicated: the best processing method of Ca~(2+) reduces the injury of flood to the hot pepper seedling is: 10mmol·L~(-1) Ca~(2+) soaking seeds processing.
The measuration's result indicated that the extraneous Ca~(2+) processing can suppress effectively to the chlorophyll destruction, membrane penetrable elevation and root activity being weaken under flood stress, and relatively enhanced the fresh weight and dry weight of seedling, reduced root system's damage. These indicated the extraneous Ca~(2+) processing could enhance the resistance of hot pepper seedling to overhead flooding injury.
The measuration about oxidation resistance enzyme activity and the non-enzyme oxidation inhibitor content demonstrate that after the Ca~(2+) processing, the hot pepper seedling leaf has the function maintain high level ROS metabolism, its mechanism possibly is advantageous to induce biosynthesis and keep activity about anti-oxidase enzyme of SOD et al., and non-enzyme oxidation inhibitor of ASA and GSH. Thus enhances the ability of plant to eliminate the free radical, reduced the membrane system's peroxidation damage. This possibly is a physiology biochemistry foundation of Ca~(2+) could enhance the resistance of hot pepper seedling to overhead flooding injury.
Exogenous Ca~(2+) application in nutrient solution could relieve the harm caused by flood stress and result in improved growth of roots, increased fresh and dry weight of pepper seedlings. Compared with the seedlings with flood alone treatment, the activities of SDH, MDH and ADH were promoted, while LDH activity was inhibited, accumulations of lactate and acetaldehyde in pepper seedling roots under exogenous application Ca~(2+) treatment. It indicated that Ca~(2+) could relieve the harm of flood stress, through regulating respiratory metabolism of roots.
Exogenous Ca~(2+) application treatment could increase the contents of and solubility sugar, cane sugar and fruit sugar in roots, stems and leaves of pepper seedlings, decreased the content of starch and glucose in pepper seedling roots, which could improve in the ratio of root carbohydrate concentration to leaves carbohydrate concentration, and promote the appearance of carbohydrate metabolism; adapting the flood stress. It indicated that Ca~(2+) could adjust carbohydrate contents and relieve the harm of flood stress.
引文
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