丝栗栲和苦槠幼苗生长状况和气体交换特征对增温和施氮的响应
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摘要
常绿阔叶林是亚热带地区的地带性植被，其重要物种对全球变化的作用与响应是常绿阔叶林植被动态研究的重要内容之一。丝栗栲（Castanopsis fargesii）和苦槠（Castanopsissclerophylla）是中亚热带典型常绿阔叶林的主要优势树种。本论文首先测定了苦槠、青冈栎和细叶青冈盆栽幼苗的光合生理指标以及丝栗栲和苦槠当年生叶片的光合日动态特征，然后以中亚热带典型常绿阔叶林主要优势种丝栗栲和苦槠幼苗为研究对象，以实验学手段为主，利用MSR-2420红外辐射增温装置，通过人工定期向土壤施加NH4NO3，实现丝栗栲和苦槠自然生长环境条件中的增温和施氮设置，采用植物生理生态学、生物化学、野外数据获取与室内模拟分析相结合的方法，利用LI-6400R便携式光合仪、OS-30P便携式荧光仪多个国际先进仪器，应用Farquhar光合作用生物化学机理模型，研究了丝栗栲和苦槠幼苗叶片气体交换相关指标对增温和施氮的响应差异及其机理。结果表明，增温、施氮及其交互作用对丝栗栲和苦槠生长状况和气体交换指标的影响程度不同。通过综合分析各项生理指标，苦槠幼苗对增温、施氮及其交互作用的适应能力更强。本研究结果可为认识未来气候变化情景下亚热带常绿阔叶林碳循环特征及其碳汇/源变化提供叶片水平的科学依据。主要结果如下：
（1）总体上，在不同的光照强度下，光合能力强弱顺序为苦槠＞青冈栎＞细叶青冈。CO2浓度的增加对三个树种光合作用具有明显的促进作用，净光合速率、水分利用率提高，光补偿点、气孔导度、蒸腾速率下降，叶片对CO2增长的这种反应对其生长是有利的，苦槠光合促进作用最明显，其次是青冈栎和细叶青冈。
（2）丝栗栲当年生叶片的光合能力显著高于苦槠栲叶片。无论是丝栗栲叶片还是苦槠栲叶片，空气相对湿度（RH）低、叶片与空气之间的水蒸气压差（VPD）大时，气孔导度（Gs）减小，气孔限制值（Ls）增大，胞间CO2浓度（Ci）降低，净光合速率（Pn）下降；中午光合有效辐射（PAR）最大，水蒸气压差（VPD）、气孔限制值（Ls）达到最大值，空气相对湿度（RH）、气孔导度（Gs）、胞间CO2浓度（Ci）达到最小值出现光合午休现象。
（3）随着施氮量的增加丝栗栲叶片和苦槠栲叶片的净光合速率（Pn）、蒸腾速率（E）、气孔导度（Gs）、表观量子效率（AQY）、表观CO2羧化效率（CCE）均有显著升高，但施氮量的增加并未显著影响叶片的水分利用率（WUE），而且同一施氮量丝栗栲叶片和苦槠栲叶片的水分利用率（WUE）之间无显著差别。随着施氮量的增加，丝栗栲叶片还是苦槠栲叶片的叶绿素a（Chl a）、叶绿素b（Chl b）、Chl a+Chl b均显著提高，这表明两种植物均对外源氮肥有一定的自我适应机制。
（4）与对照相比较，施低氮、施高氮、增温、增温和施低氮、增温和施高氮均显著提高了丝栗栲和苦槠的地径和树高。同一测量时间、同一处理条件下，苦槠的地径和树高均显著高于丝栗栲（P＜0.01），无论对照、增温、施氮与增温和施氮处理。与对照相比较，增温、施氮、增温和施氮均显著提高了苦槠总生物量、地上部分生物量和地下部分生物量。然而，与对照相比较，只有施氮引起了苦槠地上部分生物量与地下部分生物量比值的显著提高；而增温、增温和施氮与对照之间并无显著差别。
（5）较对照，增温、施氮、增温和施氮均显著提高了丝栗栲和苦槠的最大净光合速率和表观量子效率。相同处理条件下，最大净光合速率和表观量子效率均表现为苦槠＞丝栗栲，无论对照、增温、施氮、增温和施氮，丝栗栲和苦槠的暗呼吸速率并未表现出明显的大小顺序。较对照，增温、施氮、增温和施氮均显著提高了丝栗栲和苦槠的叶绿素a、叶绿素b和总叶绿素含量。在相同处理条件下，叶绿素a含量、叶绿素b含量和叶绿素总含量均表现为苦槠＞丝栗栲。
Broad-leaved evergreen forest is a zonal vegetation type. The effect and response of itsimportant species to global change is one of the key research contents in vegetationdynamics.Castanopsis fargesii and Castanopsis sclerophylla are the dominant species oftypical evergreen broad-leaved forest in mid-subtropical zone. A sequential interaction oftemperature by the artifical warming facilities MSR-2420and nitrogen by NH4NO3applicationto the soil to Castanopsis fargesii and Castanopsis sclerophylla seedlings was set up in JiangxiDagangshan State Forest Ecosystem Research Station from2009to2011. By the application ofseveral international advanced instruments and Farquhar photosynthetic biochemicalmechanism model, at the same time, through the methods of plant physiological ecology,biochemistry, data collection outside and data analysis inside, the Castanopsis fargesii andCastanopsis sclerophylla seedlings response of growth and leaf gas exchange was illustrated.The results showed that the effects of warming, nitrogen application and its interaction to theCastanopsis fargesii and Castanopsis sclerophylla seedlings was different. On the basis ofintegrated analysis of the experimental data, the adaptive ability of Castanopsis sclerophyllaseedlings to warming, nitrogen application and its interaction was higher than that ofCastanopsis sclerophylla seedlings. The results will provide the theory in the leaf level for thecarbon cycle and caron sink/source change of evergreen broad-leaved forest in mid-subtropicalzone under future climate change scenarios.The main results are as the following：
（1）Overall, in a different light intensities, the order of the photosynthetic capacity isCastanopsis sclerophylla> Cyclobalanopsis glauca（Thunb.） Oerst> Cyclobalanopsismyrsinaefolia （Blume） Oerst. The increase of CO2concentration has a significant role on thethree tree species photosynthesis, net photosynthetic rate, water use efficiency increases, lightcompensation point, stomatal conductance, transpiration rate decreases, it is favorable for thethe growth of leaf to CO2increases, which Castanopsis sclerophylla photosynthesis to promote the most significant effect, followed by Cyclobalanopsis glauca（Thunb.） Oerst andCyclobalanopsis myrsinaefolia （Blume） Oerst.
（2）The photosynthetic capacity of Castanopsis fargesii is significantly higher thanCastanopsis sclerophylla. When the low air relative humidity （RH）, high water vaporpressure between leaf and air （VPD）, stomatal conductance （Gs） is reduced, the stomatallimitation value （Ls） increases, lower intercellular CO2concentration （Ci）, netphotosynthetic rate （Pn） decreased; photosynthetically active radiation （PAR） at noon ishighest, the water vapor pressure difference （VPD）, stomatal limitation value （Ls） reachesthe maximum, relative air humidity （RH）, stomatal conductance degrees （Gs）, intercellularCO2concentration （Ci） reaches a minimum for photosynthetic midday depression.
（3）With the increase in the amount of nitrogen, Castanopsis fargesii and Castanopsissclerophylla, net photosynthetic rate （Pn）, transpiration rate （E）, stomatal conductancedegrees （Gs）, apparent quantum yield （AQY） efficiency, CO2carboxylation efficiency（CCE） have significantly increased, but the increased amount of nitrogen does notsignificantly affect leaf water use efficiency （WUE）, and the same nitrogen rate, water useefficiency （WUE） is not a significant difference. With the increase of the amount of nitrogen,chlorophyll a （Chl a）, chlorophyll b （the Chl b）, Chl a+Chl b of Castanopsis fargesii andCastanopsis sclerophylla were significantly increased, which reflects the two plants are on leafage increase in a certain degree of self-adaptive mechanisms.
（4）On the contrast, low nitrogen, high nitrogen, warming, warming and facilities withlow nitrogen, warming and applying high nitrogen significantly improved diameter and treeheight of Castanopsis fargesii and Castanopsis sclerophylla. The same measurement time, thesame processing conditions, Castanopsis sclerophylla diameter and height were significantlyhigher than Castanopsis fargesii（P <0.01）regardless of the control. On the contrast, warming,nitrogen, warming and nitrogen fertilization significantly improve Castanopsis sclerophyllatotal biomass, aboveground biomass and underground part of the biomass. However, on thecontrast, nitrogen fertilizer caused the value of the aboveground biomass and underground part of the biomass significantly improved, there was no significant difference with control ofwarming and nitrogen fertilization.
（5）Compared with the respective control, elevated warm and applied nitrogen alone andin combination significantly improved Castanopsis fargesii and Castanopsis sclerophylla,maximum net photosynthetic rate and apparent quantum efficiency respectively. Under thesame treatments, the maximum net photosynthetic rate and apparent quantum in Castanopsissclerophylla,was bigger than Castanopsis fargesii.Regardless of the control, elevated warm andapplied nitrogen alone and in combination, Castanopsis fargesii the dark respiration rate doesnot seem to did not show the apparent size of the order. the respective control, warming,nitrogen, elevated warm and applied nitrogen alone and in combination significantlyimproved the content of chlorophyll a, chlorophyll b and total chlorophyll content in theCastanopsis sclerophylla and Castanopsis fargesii. The same processing conditions,Castanopsis sclerophylla was much more than Castanopsis fargesii based on the contents ofchlorophyll a, chlorophyll b and total chlorophyll.
（1）总体上，在不同的光照强度下，光合能力强弱顺序为苦槠＞青冈栎＞细叶青冈。CO2浓度的增加对三个树种光合作用具有明显的促进作用，净光合速率、水分利用率提高，光补偿点、气孔导度、蒸腾速率下降，叶片对CO2增长的这种反应对其生长是有利的，苦槠光合促进作用最明显，其次是青冈栎和细叶青冈。
（2）丝栗栲当年生叶片的光合能力显著高于苦槠栲叶片。无论是丝栗栲叶片还是苦槠栲叶片，空气相对湿度（RH）低、叶片与空气之间的水蒸气压差（VPD）大时，气孔导度（Gs）减小，气孔限制值（Ls）增大，胞间CO2浓度（Ci）降低，净光合速率（Pn）下降；中午光合有效辐射（PAR）最大，水蒸气压差（VPD）、气孔限制值（Ls）达到最大值，空气相对湿度（RH）、气孔导度（Gs）、胞间CO2浓度（Ci）达到最小值出现光合午休现象。
（3）随着施氮量的增加丝栗栲叶片和苦槠栲叶片的净光合速率（Pn）、蒸腾速率（E）、气孔导度（Gs）、表观量子效率（AQY）、表观CO2羧化效率（CCE）均有显著升高，但施氮量的增加并未显著影响叶片的水分利用率（WUE），而且同一施氮量丝栗栲叶片和苦槠栲叶片的水分利用率（WUE）之间无显著差别。随着施氮量的增加，丝栗栲叶片还是苦槠栲叶片的叶绿素a（Chl a）、叶绿素b（Chl b）、Chl a+Chl b均显著提高，这表明两种植物均对外源氮肥有一定的自我适应机制。
（4）与对照相比较，施低氮、施高氮、增温、增温和施低氮、增温和施高氮均显著提高了丝栗栲和苦槠的地径和树高。同一测量时间、同一处理条件下，苦槠的地径和树高均显著高于丝栗栲（P＜0.01），无论对照、增温、施氮与增温和施氮处理。与对照相比较，增温、施氮、增温和施氮均显著提高了苦槠总生物量、地上部分生物量和地下部分生物量。然而，与对照相比较，只有施氮引起了苦槠地上部分生物量与地下部分生物量比值的显著提高；而增温、增温和施氮与对照之间并无显著差别。
（5）较对照，增温、施氮、增温和施氮均显著提高了丝栗栲和苦槠的最大净光合速率和表观量子效率。相同处理条件下，最大净光合速率和表观量子效率均表现为苦槠＞丝栗栲，无论对照、增温、施氮、增温和施氮，丝栗栲和苦槠的暗呼吸速率并未表现出明显的大小顺序。较对照，增温、施氮、增温和施氮均显著提高了丝栗栲和苦槠的叶绿素a、叶绿素b和总叶绿素含量。在相同处理条件下，叶绿素a含量、叶绿素b含量和叶绿素总含量均表现为苦槠＞丝栗栲。
Broad-leaved evergreen forest is a zonal vegetation type. The effect and response of itsimportant species to global change is one of the key research contents in vegetationdynamics.Castanopsis fargesii and Castanopsis sclerophylla are the dominant species oftypical evergreen broad-leaved forest in mid-subtropical zone. A sequential interaction oftemperature by the artifical warming facilities MSR-2420and nitrogen by NH4NO3applicationto the soil to Castanopsis fargesii and Castanopsis sclerophylla seedlings was set up in JiangxiDagangshan State Forest Ecosystem Research Station from2009to2011. By the application ofseveral international advanced instruments and Farquhar photosynthetic biochemicalmechanism model, at the same time, through the methods of plant physiological ecology,biochemistry, data collection outside and data analysis inside, the Castanopsis fargesii andCastanopsis sclerophylla seedlings response of growth and leaf gas exchange was illustrated.The results showed that the effects of warming, nitrogen application and its interaction to theCastanopsis fargesii and Castanopsis sclerophylla seedlings was different. On the basis ofintegrated analysis of the experimental data, the adaptive ability of Castanopsis sclerophyllaseedlings to warming, nitrogen application and its interaction was higher than that ofCastanopsis sclerophylla seedlings. The results will provide the theory in the leaf level for thecarbon cycle and caron sink/source change of evergreen broad-leaved forest in mid-subtropicalzone under future climate change scenarios.The main results are as the following：
（1）Overall, in a different light intensities, the order of the photosynthetic capacity isCastanopsis sclerophylla> Cyclobalanopsis glauca（Thunb.） Oerst> Cyclobalanopsismyrsinaefolia （Blume） Oerst. The increase of CO2concentration has a significant role on thethree tree species photosynthesis, net photosynthetic rate, water use efficiency increases, lightcompensation point, stomatal conductance, transpiration rate decreases, it is favorable for thethe growth of leaf to CO2increases, which Castanopsis sclerophylla photosynthesis to promote the most significant effect, followed by Cyclobalanopsis glauca（Thunb.） Oerst andCyclobalanopsis myrsinaefolia （Blume） Oerst.
（2）The photosynthetic capacity of Castanopsis fargesii is significantly higher thanCastanopsis sclerophylla. When the low air relative humidity （RH）, high water vaporpressure between leaf and air （VPD）, stomatal conductance （Gs） is reduced, the stomatallimitation value （Ls） increases, lower intercellular CO2concentration （Ci）, netphotosynthetic rate （Pn） decreased; photosynthetically active radiation （PAR） at noon ishighest, the water vapor pressure difference （VPD）, stomatal limitation value （Ls） reachesthe maximum, relative air humidity （RH）, stomatal conductance degrees （Gs）, intercellularCO2concentration （Ci） reaches a minimum for photosynthetic midday depression.
（3）With the increase in the amount of nitrogen, Castanopsis fargesii and Castanopsissclerophylla, net photosynthetic rate （Pn）, transpiration rate （E）, stomatal conductancedegrees （Gs）, apparent quantum yield （AQY） efficiency, CO2carboxylation efficiency（CCE） have significantly increased, but the increased amount of nitrogen does notsignificantly affect leaf water use efficiency （WUE）, and the same nitrogen rate, water useefficiency （WUE） is not a significant difference. With the increase of the amount of nitrogen,chlorophyll a （Chl a）, chlorophyll b （the Chl b）, Chl a+Chl b of Castanopsis fargesii andCastanopsis sclerophylla were significantly increased, which reflects the two plants are on leafage increase in a certain degree of self-adaptive mechanisms.
（4）On the contrast, low nitrogen, high nitrogen, warming, warming and facilities withlow nitrogen, warming and applying high nitrogen significantly improved diameter and treeheight of Castanopsis fargesii and Castanopsis sclerophylla. The same measurement time, thesame processing conditions, Castanopsis sclerophylla diameter and height were significantlyhigher than Castanopsis fargesii（P <0.01）regardless of the control. On the contrast, warming,nitrogen, warming and nitrogen fertilization significantly improve Castanopsis sclerophyllatotal biomass, aboveground biomass and underground part of the biomass. However, on thecontrast, nitrogen fertilizer caused the value of the aboveground biomass and underground part of the biomass significantly improved, there was no significant difference with control ofwarming and nitrogen fertilization.
（5）Compared with the respective control, elevated warm and applied nitrogen alone andin combination significantly improved Castanopsis fargesii and Castanopsis sclerophylla,maximum net photosynthetic rate and apparent quantum efficiency respectively. Under thesame treatments, the maximum net photosynthetic rate and apparent quantum in Castanopsissclerophylla,was bigger than Castanopsis fargesii.Regardless of the control, elevated warm andapplied nitrogen alone and in combination, Castanopsis fargesii the dark respiration rate doesnot seem to did not show the apparent size of the order. the respective control, warming,nitrogen, elevated warm and applied nitrogen alone and in combination significantlyimproved the content of chlorophyll a, chlorophyll b and total chlorophyll content in theCastanopsis sclerophylla and Castanopsis fargesii. The same processing conditions,Castanopsis sclerophylla was much more than Castanopsis fargesii based on the contents ofchlorophyll a, chlorophyll b and total chlorophyll.
引文
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张守仁.叶绿素荧光动力学参数的意义及讨论.植物学通报，1999，16（4）：444~448
张守仁，高荣孚.光胁迫下杂种杨无性系光合生理生态特性的研究.植物生态学报，2000，24（5）：528~533
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张阿宏，齐孟文，张晔晖.调制叶绿素荧光动力学参数及其计量关系的意义和公理化讨论.核农学报，2008，22（6）：909~912