长期不同施肥对玉米叶片光合作用及光系统功能的影响
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摘要
提高玉米产量是我国解决粮食安全问题的主要途径,施肥是提高玉米单产最为有效的途径,研究施肥对玉米光合机理影响意义深远。玉米的长期定位肥料试验定向培育了土壤质量和生产功能特点显著不同的农田生态系统,为研究长期不同施肥模式对土壤养分、玉米生产能力、肥料作用机理以及选择科学施肥模式提供了最优的试验平台；光合速率和光合效率测定设备,以及光响应曲线、快速叶绿素荧光诱导动力学曲线等数据分析技术为探明作物的光合性能提供了技术基础。本研究以玉米长期定位试验为平台,利用光合性能测定、光合效率测定、OJIP曲线分析以及酶活性测定等技术手段,研究长期施用有机肥、氮肥、磷肥、钾肥对玉米穗位叶叶片气体交换性能、光响应特性、光系统功能、抗衰老酶活性等的影响,并结合土壤肥力、玉米产量、生育指标、叶片养分含量进行综合分析,探索长期施肥影响作物产量的光合与衰老机理,得出如下结论：
(1)长期施用有机肥可以显著改善土壤养分状况,全面改善玉米植株生长及叶片营养状况,增产效果最显著；综合提高叶绿素含量和光合性能,提高玉米叶片对强光的利用能力,降低灌浆后期生理活性,提高PSⅠ反应中心的最大氧化还原能力,提高CAT活性。
(2)长期施用氮肥可引起土壤酸化,提高土壤有机质、氮素、有效磷钾含量,显著改善玉米生长和叶片营养状况,提高玉米经济产量；提高叶绿素含量和光合性能,提高对强光的利用能力,改善叶片生理活性,改善PSⅡ整个电子传递链的性能和受体侧电子传递能力,提高SOD、POD活性。氮肥对光合性能以及产量的影响较磷、钾肥明显,主要因为氮肥提高叶绿素含量、增强对强光的利用效率；但施氮过量可以引起较严重的光合功能损伤,导致生育后期强光抑制,造成产量降低。
(3)长期施用磷肥显著提高土壤全磷、有效磷含量,显著提高叶绿素含量,明显改善灌浆后期的光合性能,降低灌浆后期的光抑制,增强对弱光的利用能力,提高玉米的耐阴性,保持穗位叶生育后期较高的生理活性,延长有效光合持续期,提高PSⅡ电子传递链的性能,改善PSⅠ反应中心氧化还原能力,提高POD、CAT。磷肥增强对弱光的利用效率,降低叶片强光抑制,延长有效光合持续期是增产的主要原因。
(4)长期施用钾肥显著提高土壤全钾、速效钾含量,改善叶片气孔调节能力,略提高强光利用效率,引起生育后期生理活性降低,出现较严重的光抑制,提高PSⅡ电子传递链的性能,引起SOD、CAT活性下降,导致作物早熟。在高量有机肥条件下,施用钾肥对光合的促进作用不显著,甚至对部分光合性能有抑制的作用。
(5)7种模型对比研究表明,直角双曲线修正模型和二次回归模型在光饱和、非光饱和和光抑制下均能较好地对玉米灌浆期穗位叶光响应曲线进行拟合,拟合的光合参数与实测值相近,拟合的光合参数信息较多,光响应参数求解中可用两种模型求解的平均值作为最终光合参数值。
(6)随土壤养分含量增加,叶片全磷、全钾含量提高,光和性能得到改善；玉米叶片全氮含量与PSⅡ功能关系密切,而全磷、全钾与PSⅠ功能关系密切；玉米叶片叶绿素含量与光响应参数、光系统功能关系密切,而光合气体交换性能与其他光合指标间无显著相关关系；PSⅡ电子传递性能与叶片POD活性关系密切,SOD活性提高有抑制PSⅠ的氧化还原活性的作用。
Improving corn production is the main way to solve the problem of food security in China. The long-term fertilizer experiment of corn directional cultivated the markedly different soil quality and production function of characteristics in the farmland ecosystem. Therefore it provides an optimal test platform, for the study of soil nutrients, corn production capacity, fertilizer effect mechanism and selection of scientific fertilization mode under long-term different fertilization pattern. Measurement equipment for photosynthetic rate and photosynthetic efficiency, and data analysis techniques for the light response curve, rapid chlorophyll fluorescence induction kinetics curve and PS I absorption curve of820nm light, provided technical basis of proving crop photosynthetic performance.
With long-term trial corn as the platform, this study using methods such as photosynthetic performance measurement, analysis technology and enzyme activity measurement technology, researched the effect of the long-term application of organic fertilizer, nitrogen fertilizer, phosphate fertilizer, potash fertilizer, on the gas exchange parameters, light response curve, the chlorophyll fluorescence induction kinetics curve, anti-ageing enzyme activity in corn ear leaves. And made a comprehensive analysis of soil fertility and fertilizer rates, maize yield, growth indexes, and leaf nutrient content, to explore the mechanism of photosynthesis and aging for changing crop yield under long-term fertilization. The conclusions is following:
(1) Long-term application of organic fertilizer could significantly improve the soil nutrient status, improve plant growth status and leaf nutritional status of maize, and the effect of production is the most significant. It could increase the chlorophyll content and photosynthetic performance, improve the highlight utilization in maize leaf, reduce physiological activity in late grouting, improve the biggest REDOX ability in PS I reaction center, improve the activity of CAT.
(2) Long-term application of nitrogen fertilizer could cause soil acidification, increase soil organic matter, nitrogen, phosphorus, potassium content, improve maize growth and leaf nutrition significantly, improve the corn output, increase the chlorophyll content and photosynthetic performance, improve their ability to the use of highlight, and improve the leaf physiological activity, improve the performance of the electron transport chain in PS II, improve the activity of SOD and POD. The effect of Nitrogen fertilizer on photosynthetic characteristics and yield was more obvious than p and k fertilizers, it was mainly because of nitrogenous fertilizer can increase chlorophyll content and the utilization efficiency of highlight. But excessive nitrogen could cause serious injure of photosynthetic function, cause strong inhibition in late grouting, reduce the production.
(3) Long-term use of phosphate fertilizer could improve the soil total phosphorus and available phosphorus content significantly, increase the chlorophyll content, improve the photosynthetic performance obviously in late grouting, reduce the photoinhibition in late grouting, increase the use of weak light, improve Shade-endurance of corn to negative, keep the ear leaf higher physiological activity, extend the effective photosynthetic duration, improve the performance of the electron transport chain in PS II, improve the REDOX ability of PS I reaction center, improve the activity of POD, CAT. The main reason for the increase production of phosphate was that it could strengthen the utilization efficiency of weak light, reduce strong light inhibition, extend the effective photosynthetic duration.
(4) Long-term use of K fertilizer could increase soil total potassium, available potassium content significantly, improve the Pn, reduce the Gs, Ci, Tr, improve the stomata adjustment ability, improve the utilization efficiency of highlight, due to physiological activity decreased in late grouting, appear serious photoinhibition, improve the performance of the electron transfer chain in PS II and the electron transfer ability on the receptor side, decline the activity of SOD, CAT, lead to crops premature aging (precocity). Under the condition of high amount of organic fertilizer, the effect of K fertilizer on promoting photosynthetic was not significant, even K fertilizer could inhibit part of photosynthetic performance.
(5) The comparative study on seven kinds of model contrast research showed that the right Angle hyperbolic correction model and quadratic regression model could fit corn light response curve preferably in the light saturation, light unsaturation and light inhibition. The fitting photosynthetic parameters were more, and the photosynthetic parameters of fitting were close to the measured values. And using the average of the two kinds of models as the final solution of photosynthesis parameter values.
(6) With the soil nutrient content increasing, the maize growth and yield improved, total phosphorus, total potassium content increased. The corn leaf total nitrogen content is closely associated with PSⅡ function, and the relationship between total phosphorus, potassium and PS I function close. The relationship among the corn leaf chlorophyll content and optical response parameters, photosystem function and decay resistance, but has no obvious relation with photosynthetic gas exchange parameters. There was no significant correlation between the photosystem function, anti-aging performance and apparent photosynthesis. The electron transfer performance in PS II was close to leaf POD activity, and the improving of SOD activity could inhibit the action of the PS I activity.
(1)长期施用有机肥可以显著改善土壤养分状况,全面改善玉米植株生长及叶片营养状况,增产效果最显著；综合提高叶绿素含量和光合性能,提高玉米叶片对强光的利用能力,降低灌浆后期生理活性,提高PSⅠ反应中心的最大氧化还原能力,提高CAT活性。
(2)长期施用氮肥可引起土壤酸化,提高土壤有机质、氮素、有效磷钾含量,显著改善玉米生长和叶片营养状况,提高玉米经济产量；提高叶绿素含量和光合性能,提高对强光的利用能力,改善叶片生理活性,改善PSⅡ整个电子传递链的性能和受体侧电子传递能力,提高SOD、POD活性。氮肥对光合性能以及产量的影响较磷、钾肥明显,主要因为氮肥提高叶绿素含量、增强对强光的利用效率；但施氮过量可以引起较严重的光合功能损伤,导致生育后期强光抑制,造成产量降低。
(3)长期施用磷肥显著提高土壤全磷、有效磷含量,显著提高叶绿素含量,明显改善灌浆后期的光合性能,降低灌浆后期的光抑制,增强对弱光的利用能力,提高玉米的耐阴性,保持穗位叶生育后期较高的生理活性,延长有效光合持续期,提高PSⅡ电子传递链的性能,改善PSⅠ反应中心氧化还原能力,提高POD、CAT。磷肥增强对弱光的利用效率,降低叶片强光抑制,延长有效光合持续期是增产的主要原因。
(4)长期施用钾肥显著提高土壤全钾、速效钾含量,改善叶片气孔调节能力,略提高强光利用效率,引起生育后期生理活性降低,出现较严重的光抑制,提高PSⅡ电子传递链的性能,引起SOD、CAT活性下降,导致作物早熟。在高量有机肥条件下,施用钾肥对光合的促进作用不显著,甚至对部分光合性能有抑制的作用。
(5)7种模型对比研究表明,直角双曲线修正模型和二次回归模型在光饱和、非光饱和和光抑制下均能较好地对玉米灌浆期穗位叶光响应曲线进行拟合,拟合的光合参数与实测值相近,拟合的光合参数信息较多,光响应参数求解中可用两种模型求解的平均值作为最终光合参数值。
(6)随土壤养分含量增加,叶片全磷、全钾含量提高,光和性能得到改善；玉米叶片全氮含量与PSⅡ功能关系密切,而全磷、全钾与PSⅠ功能关系密切；玉米叶片叶绿素含量与光响应参数、光系统功能关系密切,而光合气体交换性能与其他光合指标间无显著相关关系；PSⅡ电子传递性能与叶片POD活性关系密切,SOD活性提高有抑制PSⅠ的氧化还原活性的作用。
Improving corn production is the main way to solve the problem of food security in China. The long-term fertilizer experiment of corn directional cultivated the markedly different soil quality and production function of characteristics in the farmland ecosystem. Therefore it provides an optimal test platform, for the study of soil nutrients, corn production capacity, fertilizer effect mechanism and selection of scientific fertilization mode under long-term different fertilization pattern. Measurement equipment for photosynthetic rate and photosynthetic efficiency, and data analysis techniques for the light response curve, rapid chlorophyll fluorescence induction kinetics curve and PS I absorption curve of820nm light, provided technical basis of proving crop photosynthetic performance.
With long-term trial corn as the platform, this study using methods such as photosynthetic performance measurement, analysis technology and enzyme activity measurement technology, researched the effect of the long-term application of organic fertilizer, nitrogen fertilizer, phosphate fertilizer, potash fertilizer, on the gas exchange parameters, light response curve, the chlorophyll fluorescence induction kinetics curve, anti-ageing enzyme activity in corn ear leaves. And made a comprehensive analysis of soil fertility and fertilizer rates, maize yield, growth indexes, and leaf nutrient content, to explore the mechanism of photosynthesis and aging for changing crop yield under long-term fertilization. The conclusions is following:
(1) Long-term application of organic fertilizer could significantly improve the soil nutrient status, improve plant growth status and leaf nutritional status of maize, and the effect of production is the most significant. It could increase the chlorophyll content and photosynthetic performance, improve the highlight utilization in maize leaf, reduce physiological activity in late grouting, improve the biggest REDOX ability in PS I reaction center, improve the activity of CAT.
(2) Long-term application of nitrogen fertilizer could cause soil acidification, increase soil organic matter, nitrogen, phosphorus, potassium content, improve maize growth and leaf nutrition significantly, improve the corn output, increase the chlorophyll content and photosynthetic performance, improve their ability to the use of highlight, and improve the leaf physiological activity, improve the performance of the electron transport chain in PS II, improve the activity of SOD and POD. The effect of Nitrogen fertilizer on photosynthetic characteristics and yield was more obvious than p and k fertilizers, it was mainly because of nitrogenous fertilizer can increase chlorophyll content and the utilization efficiency of highlight. But excessive nitrogen could cause serious injure of photosynthetic function, cause strong inhibition in late grouting, reduce the production.
(3) Long-term use of phosphate fertilizer could improve the soil total phosphorus and available phosphorus content significantly, increase the chlorophyll content, improve the photosynthetic performance obviously in late grouting, reduce the photoinhibition in late grouting, increase the use of weak light, improve Shade-endurance of corn to negative, keep the ear leaf higher physiological activity, extend the effective photosynthetic duration, improve the performance of the electron transport chain in PS II, improve the REDOX ability of PS I reaction center, improve the activity of POD, CAT. The main reason for the increase production of phosphate was that it could strengthen the utilization efficiency of weak light, reduce strong light inhibition, extend the effective photosynthetic duration.
(4) Long-term use of K fertilizer could increase soil total potassium, available potassium content significantly, improve the Pn, reduce the Gs, Ci, Tr, improve the stomata adjustment ability, improve the utilization efficiency of highlight, due to physiological activity decreased in late grouting, appear serious photoinhibition, improve the performance of the electron transfer chain in PS II and the electron transfer ability on the receptor side, decline the activity of SOD, CAT, lead to crops premature aging (precocity). Under the condition of high amount of organic fertilizer, the effect of K fertilizer on promoting photosynthetic was not significant, even K fertilizer could inhibit part of photosynthetic performance.
(5) The comparative study on seven kinds of model contrast research showed that the right Angle hyperbolic correction model and quadratic regression model could fit corn light response curve preferably in the light saturation, light unsaturation and light inhibition. The fitting photosynthetic parameters were more, and the photosynthetic parameters of fitting were close to the measured values. And using the average of the two kinds of models as the final solution of photosynthesis parameter values.
(6) With the soil nutrient content increasing, the maize growth and yield improved, total phosphorus, total potassium content increased. The corn leaf total nitrogen content is closely associated with PSⅡ function, and the relationship between total phosphorus, potassium and PS I function close. The relationship among the corn leaf chlorophyll content and optical response parameters, photosystem function and decay resistance, but has no obvious relation with photosynthetic gas exchange parameters. There was no significant correlation between the photosystem function, anti-aging performance and apparent photosynthesis. The electron transfer performance in PS II was close to leaf POD activity, and the improving of SOD activity could inhibit the action of the PS I activity.
引文
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