大气二氧化碳浓度升高对玉米幼苗碳氮资源分配的影响
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摘要
大气CO_2浓度不断升高是全球气候变化的主要组成部分，其对个体植株生理生态过程的调节直接影响到群落组成与大气-植物-土壤圈的物质循环。在降雨量小、蒸发量大且土壤贫瘠的黄土高原，作物生长遭受着土壤干旱-复水（有限降雨）-干旱的循环过程，干旱与氮素缺乏是作物生长的主要限制因素。目前关于CO_2浓度升高对C_3植物生理过程的影响与调控的工作较多，而C4植物对CO_2浓度升高的生理生态适应性仍存在不少争论。光合调节是植株适应气候变化最直接的表现方式，体内碳氮分配与转运策略是评估植株长期适应能力的主要内容。研究CO_2浓度升高、干旱与氮素胁迫三者的交互作用对C4植物光合能力与体内碳氮分配与转运策略的影响有利于为探索旱区植物对未来气候变化的适应性提供理论依据。
本研究以水培玉米幼苗为试验材料，采用人工气候室内控制CO_2浓度，聚乙二醇（PEG-6000）模拟水分胁迫的方法，利用碳氮化学计量技术、同位素示踪技术、叶绿素荧光和气体交换测定技术，从玉米生长、光合生理、碳氮分配及转运、碳氮利用等角度，研究了水分胁迫下高浓度CO_2减轻叶片光合抑制的作用机制、源库器官活力对植株生长的影响，分析了高浓度CO_2提高源库器官活力的途径，探讨了氮胁迫抑制高浓度CO_2作用的机理，以及受旱植株对不同程度复水的生理适应能力。主要研究结果如下：
（1） CO_2倍增显著减少了受旱玉米叶片有活性的PSII反应中心数目，提高了单位反应中心光能捕获、转化与传递能力，进而增强叶片单位横截面积的能量流动效率及电子传递速率；在碳反应过程中，CO_2倍增使受旱植株具有较高饱和光合速率，表明CO_2倍增可以通过提高光能利用为碳反应提高较多还原力。
（2）在正常供氮条件下，CO_2倍增可通过增强玉米叶片PEPC的羧化能力（V_(pmax)）并减轻气孔限制（SL）缓解由水分胁迫产生的光合限制；在氮素限制条件下，CO_2倍增可通过提高玉米叶片Rubisco羧化能力（V_(max)）并减轻气孔限制，部分地减缓光合限制。CO_2倍增下氮素限制使植株向叶片分配较多氮素，提高光合氮利用效率并降低整株氮利用效率。CO_2倍增可以减轻干旱对叶片光合作用的抑制，但随着植物氮素供应水平不同，其受抑程度也存在一定差异。
（3）CO_2倍增加快了受旱植株源叶碳输出速度、老叶碳输出量、以及库叶碳输入量，并减少了碳素从地下部向地上部的反向运输。CO_2倍增延长了新固定氮素在根系中的存留时间，增加了库叶氮素的总输入量。CO_2倍增改变了体内碳氮分配与再利用方式，使库叶碳氮素供应显著增加，并增强了植株生物量与碳氮素积累，表明CO_2倍增有助于减轻植株受水分胁迫的影响。
（4）CO_2浓度升高显著提高了玉米整株氮素积累量与库叶氮积累量。停止氮素供应后，高浓度CO_2下水分胁迫处理通过减缓玉米功能叶“新氮”输入速度，加快库叶“新氮”输入速度，并减少库叶“老氮”输入量以维持受旱植株生长，对老叶氮素的输出没有显著提高。
（5）受旱玉米复水后，CO_2倍增较正常CO_2使玉米植株具有较低的含水量与较高的生长速率，并具有较高的光合能力(F_v/F_m, Φ_(PSII), P_n, P_n/T_rand P_n/G_s)与新叶生长潜力，即使在低氮条件下也是如此，说明CO_2倍增通过较高的光合能力与较低含水量使植株提高水分利用效率、增强生物量积累。
Rises in ambient CO_2are expected to cause global climate changes, includingincreases in air temperature and shifts of regional scale rainfall patterns, which leadto decreased soil water availability in some areas of the world. Elevated CO_2affectplant physiological and ecosystem processes and probably lead to the suppression ofplant N availability that limits the effect of CO_2fertilization. Previous study haveshown that C_3plants under elevated CO_2often maintain growth during short termdrought due to improved water use efficiency, however, reduce long-term adaptionand result in down-regulation of photosynthesis. The maintenance of rapid growthunder conditions of CO_2enrichment is directly related to the capacity ofphotosynthesis and carbon and nitrogen transport in plants and its contribution to thenew foliar formation. Less is known about the phorosynthesis and carbon andnitrogen transport in C4plants in response to drought, N limitation, precipitationfrequency and increasing CO_2.
To test the effects of water and nitrogen limitation on plants under elevated CO_2,maize (Zea mays), the world's most important C4crop, was planted to experiencecombined elevated CO_2(380or750μmolmol~(-1), climate chamber), water stress (15%PEG-6000) and nitrogen limitation (N deficiency treated since the144th droughthour) and rewatered at three intensities (300mL,600mL,900mL of distilled water).During the growing period, the performance of PSII and electron transport,stomatallimitation, non-stomatal limitation, photosynthetic potential parameters, leafnitrogen use efficiency, patterns of carbon and nitrogen delivery, and new leafproductivities of the maize plants were investigated using chlorophyll-a fluorescenceOJIP induction curves, A/Cicurves and~(13)C and~(15)N as tracers.
(1). Compared to water-stressed maize under atmospheric CO_2, the elevated CO_2treatment interacted with water stress decreased number of active reactioncenters but increased antenna size and energy flux (absorb photon flux, trapping fluxand electron transport flux) of per reaction center in PSII. So the electron transportrate (J) was increased, despite of the indistinctively changed quantum yield forelectron transport and energy dissipation. In carbon reaction, the combination ofelevated CO_2and water stress treatment had the robust saturated photosynthetic rate(A_(sat)). This study demonstrates that maize at doubled CO_2was capable oftransporting more electron flow into carbon reaction.
(2). Elevated CO_2could alleviate drought-induced photosynthetic limitationthrough increasing capacity of PEPC carboxylation (Vpmax) and decreasing stomatallimitations (SL). The N deficiency exacerbated drought-induced photosynthesislimitations in ambient CO_2. Elevated CO_2partially alleviated the limitation inducedby drought and N deficiency through improving the capacity of Rubiscocarboxylation (Vmax) and decreasing SL. Plants with N deficiency transported moreN to their leaves at elevated CO_2, leading to a high photosynthetic nitrogen-useefficiency but low whole-plant nitrogen-use efficiency. The stress mitigation byelevated CO_2under N deficiency conditions was not enough to improving plant Nuse efficiency and biomass accumulation. The study demonstrated that elevated CO_2could alleviate drought-induced photosynthesis limitation, but the alleviation variedwith N supplies.
(3). Compared to water-stressed maize under atmospheric CO_2, the treatmentcombining elevated CO_2with water stress increased the accumulation of biomassand partitioned more carbon and nitrogen to the formation of new leaves. Maizeseedlings enhanced the carbon resource in aging leaves and the carbon pool in newleaves but decreased the carbon counterflow capability of roots. The seedlings alsohad increased residence times of new nitrogen in roots and then delivered morenitrogen to new leaves. Thus maize supported the development of new leaves atelevated levels of CO_2by altering the transport and remobilization of carbon andnitrogen. In drought presence condition, increased activity of new leaves to storecarbon and nitrogen sustains enhanced growth under elevated CO_2in maize.
(4). Elevated CO_2significently increased the accumulation of nitrogen inwhole plant and new leaves. With nitrogen starvation, elevated CO_2retard the newnitrogen transport in functional leaves, accelerate the new nitrogen transport in new leaves to sustains growth under drought.
(5). After they were rewatered, pre-drought stressed and N limited plants withambient CO_2increased their water content higher than that of elevated CO_2, whilethe enhancement of growth rate were negatively proportional to the increasing plantwater content. Elevated CO_2could help rewatered seedlings to have higherphotosynthetic capacity (F_v/F_m, Φ_(PSII), P_n, P_n/T_rand P_n/G_s) and new leaf recoveryability under low water content, no matter the seedlings suffered nitrogen deficiencyor not. The study demonstrated that elevated CO_2could help drought stressedseedlings to have higher carbon assimilation rates under low water uptakes, as aresult to improve leaf water use efficiency, which allows the plants to have muchbetter performance under drought following being re-watered.
本研究以水培玉米幼苗为试验材料，采用人工气候室内控制CO_2浓度，聚乙二醇（PEG-6000）模拟水分胁迫的方法，利用碳氮化学计量技术、同位素示踪技术、叶绿素荧光和气体交换测定技术，从玉米生长、光合生理、碳氮分配及转运、碳氮利用等角度，研究了水分胁迫下高浓度CO_2减轻叶片光合抑制的作用机制、源库器官活力对植株生长的影响，分析了高浓度CO_2提高源库器官活力的途径，探讨了氮胁迫抑制高浓度CO_2作用的机理，以及受旱植株对不同程度复水的生理适应能力。主要研究结果如下：
（1） CO_2倍增显著减少了受旱玉米叶片有活性的PSII反应中心数目，提高了单位反应中心光能捕获、转化与传递能力，进而增强叶片单位横截面积的能量流动效率及电子传递速率；在碳反应过程中，CO_2倍增使受旱植株具有较高饱和光合速率，表明CO_2倍增可以通过提高光能利用为碳反应提高较多还原力。
（2）在正常供氮条件下，CO_2倍增可通过增强玉米叶片PEPC的羧化能力（V_(pmax)）并减轻气孔限制（SL）缓解由水分胁迫产生的光合限制；在氮素限制条件下，CO_2倍增可通过提高玉米叶片Rubisco羧化能力（V_(max)）并减轻气孔限制，部分地减缓光合限制。CO_2倍增下氮素限制使植株向叶片分配较多氮素，提高光合氮利用效率并降低整株氮利用效率。CO_2倍增可以减轻干旱对叶片光合作用的抑制，但随着植物氮素供应水平不同，其受抑程度也存在一定差异。
（3）CO_2倍增加快了受旱植株源叶碳输出速度、老叶碳输出量、以及库叶碳输入量，并减少了碳素从地下部向地上部的反向运输。CO_2倍增延长了新固定氮素在根系中的存留时间，增加了库叶氮素的总输入量。CO_2倍增改变了体内碳氮分配与再利用方式，使库叶碳氮素供应显著增加，并增强了植株生物量与碳氮素积累，表明CO_2倍增有助于减轻植株受水分胁迫的影响。
（4）CO_2浓度升高显著提高了玉米整株氮素积累量与库叶氮积累量。停止氮素供应后，高浓度CO_2下水分胁迫处理通过减缓玉米功能叶“新氮”输入速度，加快库叶“新氮”输入速度，并减少库叶“老氮”输入量以维持受旱植株生长，对老叶氮素的输出没有显著提高。
（5）受旱玉米复水后，CO_2倍增较正常CO_2使玉米植株具有较低的含水量与较高的生长速率，并具有较高的光合能力(F_v/F_m, Φ_(PSII), P_n, P_n/T_rand P_n/G_s)与新叶生长潜力，即使在低氮条件下也是如此，说明CO_2倍增通过较高的光合能力与较低含水量使植株提高水分利用效率、增强生物量积累。
Rises in ambient CO_2are expected to cause global climate changes, includingincreases in air temperature and shifts of regional scale rainfall patterns, which leadto decreased soil water availability in some areas of the world. Elevated CO_2affectplant physiological and ecosystem processes and probably lead to the suppression ofplant N availability that limits the effect of CO_2fertilization. Previous study haveshown that C_3plants under elevated CO_2often maintain growth during short termdrought due to improved water use efficiency, however, reduce long-term adaptionand result in down-regulation of photosynthesis. The maintenance of rapid growthunder conditions of CO_2enrichment is directly related to the capacity ofphotosynthesis and carbon and nitrogen transport in plants and its contribution to thenew foliar formation. Less is known about the phorosynthesis and carbon andnitrogen transport in C4plants in response to drought, N limitation, precipitationfrequency and increasing CO_2.
To test the effects of water and nitrogen limitation on plants under elevated CO_2,maize (Zea mays), the world's most important C4crop, was planted to experiencecombined elevated CO_2(380or750μmolmol~(-1), climate chamber), water stress (15%PEG-6000) and nitrogen limitation (N deficiency treated since the144th droughthour) and rewatered at three intensities (300mL,600mL,900mL of distilled water).During the growing period, the performance of PSII and electron transport,stomatallimitation, non-stomatal limitation, photosynthetic potential parameters, leafnitrogen use efficiency, patterns of carbon and nitrogen delivery, and new leafproductivities of the maize plants were investigated using chlorophyll-a fluorescenceOJIP induction curves, A/Cicurves and~(13)C and~(15)N as tracers.
(1). Compared to water-stressed maize under atmospheric CO_2, the elevated CO_2treatment interacted with water stress decreased number of active reactioncenters but increased antenna size and energy flux (absorb photon flux, trapping fluxand electron transport flux) of per reaction center in PSII. So the electron transportrate (J) was increased, despite of the indistinctively changed quantum yield forelectron transport and energy dissipation. In carbon reaction, the combination ofelevated CO_2and water stress treatment had the robust saturated photosynthetic rate(A_(sat)). This study demonstrates that maize at doubled CO_2was capable oftransporting more electron flow into carbon reaction.
(2). Elevated CO_2could alleviate drought-induced photosynthetic limitationthrough increasing capacity of PEPC carboxylation (Vpmax) and decreasing stomatallimitations (SL). The N deficiency exacerbated drought-induced photosynthesislimitations in ambient CO_2. Elevated CO_2partially alleviated the limitation inducedby drought and N deficiency through improving the capacity of Rubiscocarboxylation (Vmax) and decreasing SL. Plants with N deficiency transported moreN to their leaves at elevated CO_2, leading to a high photosynthetic nitrogen-useefficiency but low whole-plant nitrogen-use efficiency. The stress mitigation byelevated CO_2under N deficiency conditions was not enough to improving plant Nuse efficiency and biomass accumulation. The study demonstrated that elevated CO_2could alleviate drought-induced photosynthesis limitation, but the alleviation variedwith N supplies.
(3). Compared to water-stressed maize under atmospheric CO_2, the treatmentcombining elevated CO_2with water stress increased the accumulation of biomassand partitioned more carbon and nitrogen to the formation of new leaves. Maizeseedlings enhanced the carbon resource in aging leaves and the carbon pool in newleaves but decreased the carbon counterflow capability of roots. The seedlings alsohad increased residence times of new nitrogen in roots and then delivered morenitrogen to new leaves. Thus maize supported the development of new leaves atelevated levels of CO_2by altering the transport and remobilization of carbon andnitrogen. In drought presence condition, increased activity of new leaves to storecarbon and nitrogen sustains enhanced growth under elevated CO_2in maize.
(4). Elevated CO_2significently increased the accumulation of nitrogen inwhole plant and new leaves. With nitrogen starvation, elevated CO_2retard the newnitrogen transport in functional leaves, accelerate the new nitrogen transport in new leaves to sustains growth under drought.
(5). After they were rewatered, pre-drought stressed and N limited plants withambient CO_2increased their water content higher than that of elevated CO_2, whilethe enhancement of growth rate were negatively proportional to the increasing plantwater content. Elevated CO_2could help rewatered seedlings to have higherphotosynthetic capacity (F_v/F_m, Φ_(PSII), P_n, P_n/T_rand P_n/G_s) and new leaf recoveryability under low water content, no matter the seedlings suffered nitrogen deficiencyor not. The study demonstrated that elevated CO_2could help drought stressedseedlings to have higher carbon assimilation rates under low water uptakes, as aresult to improve leaf water use efficiency, which allows the plants to have muchbetter performance under drought following being re-watered.
引文
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