三江平原小叶章湿地碳特征对模拟CO_2升高和氮沉降的响应
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摘要
全球CO_2浓度升高和氮沉降呈现日益增加的趋势，这些因素必将影响到植物生理生态特性，进而影响到陆地生态系统结构和功能。长期模拟大气CO_2浓度升高和氮沉降实验对预测未来全球气候变化条件下湿地生态系统生产力和碳汇功能具有十分重要的意义。本研究在我国东北三江平原湿地设置了模拟CO_2浓度升高和氮沉降的原位控制实验，系统地探讨了CO_2浓度升高和氮沉降对三江湿地优势种小叶章光合特性、光合产物和矿质元素含量的影响；以及小叶章湿地碳、水通量以及土壤呼吸的响应过程，为湿地对全球变化的适应过程提供理论依据。本研究通过OTC开顶式气室设置对照CO_2浓度（AC）、高CO_2浓度处理1（550ppm，EC1）和高CO_2浓度处理2（700ppm，EC2）3个水平；在每个气室内通过施用NH4NO3模拟3个氮沉降水平，分别为对照（0g N/m~2.a，NN）、低氮处理（4g N/m~2.a，LN）和高氮处理（8g N/m~2.a，HN）3个水平。共3次重复。获得主要结论如下：
（1）CO_2（550ppm和700ppm）升高条件下，小叶章出现光合下调现象，施氮减缓了光合下调，高氮处理下小叶章无明显的光合适应现象。在实验设计的氮沉降范围内，CO_2浓度升高降低了小叶章叶片气孔导度（Gs）、蒸腾速率（Tr），促进了水分利用效率（WUE）的提高。CO_2浓度升高显著影响了小叶章最大净光合速率（Amax）、光补偿点（LCP）、光饱和点（LSP）、最大羧化速率（Vc,max）和CO_2补偿点（г*）。光合参数对CO_2浓度升高的响应依赖于供氮水平，未施氮和低氮条件下，CO_2升高降低了小叶章叶片Amax、Vc,max，提高了LCP、LSP和г*；在高氮条件下，CO_2浓度升高未显著影响光合参数Amax、暗呼吸速率（Rd）、表观量子产量（AQE）、LCP、LSP、Vc,max和最大电子速率（Jmax）。施氮显著增加了Amax和AQE，降低了Rd、LCP和LSP，促进了小叶章光合能力的提高。CO_2浓度升高显著影响了小叶章叶片光合色素含量。未施氮和低氮条件下，CO_2浓度升高降低了小叶章叶片叶绿素a、叶绿素b、类胡萝卜素和叶绿素总量；但在高氮条件下，小叶章光合色素含量则未发生显著变化。施氮处理显著增加了小叶章叶片光合色素含量。上述结果表明，充足的氮供应可减轻甚至抑制小叶章对CO_2浓度升高的适应。小叶章出现光合适应与Vc,max的降低有关，暗示小叶章发生光合适应很大程度上是由于1,5-二磷酸核酮糖（RuBP）羧化限制。
（2）CO_2浓度升高显著增加了小叶章根中淀粉含量、各器官可溶性糖含量，但对小叶章叶片、茎的淀粉含量的影响不显著。CO_2浓度升高对小叶章叶片可溶性蛋白含量依据氮处理而异，在未施氮条件下，CO_2升高降低了可溶性蛋白含量；在低氮和高氮水平下，则增加了可溶性蛋白含量。施氮处理显著增加了小叶章茎、根中淀粉含量和总淀粉含量、小叶章根、茎和器官总可溶性糖含量以及小叶章叶片可溶性蛋白含量；对叶片中淀粉和可溶性糖含量影响不显著。
不同种类矿质元素对CO_2浓度升高的响应不同。CO_2浓度升高未对小叶章叶片C含量产生显著影响；对小叶章叶片N含量影响因氮处理不同而异。在未施氮和低氮处理下，CO_2升高降低了小叶章叶片N素含量，在高氮处理条件下，小叶章叶片N含量变化不大。CO_2浓度升高显著增加了小叶章叶片P和K含量；显著改变了小叶章C/N、C/P、N/P和N/K。施氮处理显著增加了小叶章叶片N含量；但不同程度降低了小叶章叶片P和K含量，增加了小叶章N/P和N/K。表明CO_2浓度升高和施氮改变了植物养分格局，暗示未来CO_2升高将有利于湿地植物减轻P、K元素限制。
（3）在2010-2011两个生长季中，小叶章湿地净生态系统CO_2交换量（NEE）、生态系统呼吸量（ER）和总生态系统生产力（GEP）呈现夏季高而春秋两季低的季节动态，而且有显著的年际动态，这与生长季早期大气温度、土壤温度和植被盖度有关。CO_2浓度升高降低了NEE、生态系统水分蒸发蒸腾损失量（ET），但增加了ER、GEP和生态系统水分利用效率（WUE）。NEE的降低主要是由于CO_2升高导致ER增加的幅度高于GEP增加的幅度。施氮不仅增加了生态系统碳通量，而且也削弱了CO_2浓度升高对NEE的负面效应。我们的研究结果表明，在未来伴随氮沉降逐渐增加的趋势下，CO_2浓度升高将有利于三江平原小叶章湿地碳固定。
（4）土壤呼吸的季节动态呈现夏季高的单峰曲线模式，主要归因于夏季较高的大气温度促进了植物生长和土壤微生物活性。本研究中，土壤呼吸速率和土壤温度呈现指数相关。CO_2浓度升高条件下，土壤呼吸呈现增加的趋势。施氮处理第一年土壤呼吸显著增加，而第二年土壤呼吸呈现降低的趋势，这表明施氮对土壤呼吸的促进作用随着时间的推移减弱甚至产生抑制作用。CO_2浓度升高和氮沉降对土壤呼吸的影响有显著的互作效应，二者互作显著增加了土壤呼吸，增加幅度在20%-29%。表明在未来CO_2浓度升高条件下，不管氮沉降水平如何，都将增加三江平原小叶章湿地土壤呼吸和土壤碳循环。
The increasingly increases of atmospheric CO_2and nitrogen deposition can influenceplant photosynthetic capacity and then affect ecosystem structure and functions. Thelong-term continuous simulation of atmospheric CO_2and nitrogen (N) depositionexperiments is significant to predict the future global climate change on wetland ecosystemproductivity and carbon sequestration. This study is based on the results through thecontrolled situ-experiment on the Sanjiang Plain wetland in northeast China. This studyfocus on the photosynthetic characteristics, photosynthetic product and mineral elements ofthe dominant species Calamagrostis angustifolia, and ecosystem carbon, water fluxes andsoil respiration of C.angustifolia wetland under different CO_2and N level, which provide atheoretical basis for the wetland adaptation process to global change. There were three CO_2concentration accomplished using Open Top Chambers (370ppm AC,550ppm EC1and700ppmEC2), and three N deposition level (0g N/m~2.a, NN;4g N/m~2.a, LN;8g N/m~2.a, HN) suppliedas NH4NO3. Each treatment had three replicates. The main conclusions are as follows:
(l) Photosynthetic acclimation phenomena occured when C.angustifolia exposed toelevated CO_2(550ppm and700ppm). Nitrogen addition weakened photosyntheticacclimation phenomena and photosynthetic acclimation phenomena were not found underhigh nitrogen addition. The stomatal conductance (Gs) and transpiration rate (Tr) weredecreased and the water use efficiency (WUE) was enhanced under elevated CO_2. ElevatedCO_2significantly influnced the maximum net photosynthetic rate (Amax), lightcompensation point (LCP), light saturation point (LSP), maximum carboxylation rate(Vc,max) and CO_2compensation point (г*). The response of photosynthetic parameters toelevated CO_2varied according to the level of N. Elevated CO_2decreased Amaxand Vc,max,increased LCP, LSP and г*under NN and LN treatment. However, elevated CO_2could notsignificantly affect Amax, dark respiration rate (Rd), apparent quantum efficiency (AQE),LCP, LSP, Vc,maxand maximum electron transport rate (Jmax) under HN. At the same time,N-addition enhanced Amaxand AQE, but decreased Rd,LCP and LSP, which showed that Naddition could increased photosynthetic capacity.
Elevated CO_2significantly altered the photosynthetic pigment content of leaves inC.angustifolia. The response of photosynthetic pigment content to elevated CO_2variedaccording to the level of N. The chlorophyll a, chlorophyll b, carotenoids and totalchlorophyll content of C.angustifolia were decreased under NN and LN treatment.However, the photosynthetic pigments contents of C.angustifolia grown under elevatedCO_2were not significantly lower than those grown under ambient CO_2under HN treatment.N addition enhanced chlorophyll a, chlorophyll b, carotenoids and total chlorophyll contentof C.angustifolia.
The results indicate that the photosynthetic capacity of C.angustifolia declined underlong-term elevated CO_2, but enough N addition could alleviate photosynthetic acclimation.The photosynthetic acclimation occured accompanying with significant decrease of Vc,max.Our results imply that photosynthetic acclimation under elevated CO_2is largely due toRuBP carboxylation limitation.
(2) Elevated CO_2significantly increased the starch content of root, the soluble sugarcontent of leaves, stem, root and total organ in C.angusitifoila, but had no significanteffects on the starch content of leaves and stem. The response of soluble protein content toelevated CO_2varied according to the level of nitrogen. Elevated CO_2decreased solubleprotein content of leaves in C.angusitifoila under NN treatment, but increased solubleprotein content under LN and HN treatment. N addition enhanced the starch content andsoluble sugar content of root, stem and total organ, and the soluble protein content ofleaves, but had no significant effect on the starch content and soluble sugar content ofleaves in C.angusitifoila.
The response of the mineral element to elevated CO_2varied according to the elementsspecies and the level of N. Elevated CO_2had no significant effects on the organic carbon(C) of leaves in C.angusitifoila. The response of N content of leaves to elevated CO_2variedaccording to the level of nitrogen. Elevated CO_2significantly decreased N content ofleaves in C.angusitifoila under NN and LN level but not under HN treatment. Phosphorus(P) and potassium (K) content of leaves was significantly increased under elevated CO_2.Elevated CO_2significantly influnced C/N, C/P, N/P and N/K of leaves in C.angusitifoila. Naddition significantly increased the content of N, decreased the content of P and K, andincreased N/P and N/K of leaves in C.angusitifoila. Our results show that elevated CO_2andnitrogen addition could change the nutrition pattern of plant and imply that elevated CO_2could facilitate wetland plant to mitigate P and K limitation.
(3) The seasonal dynamics of ecosystem carbon fluxes (NEE, ER and GEP) inC.angusitifoila wetland were high in mid-summer and low in the early and late growingseasons in2010and2011. In addition, there were remarkable interannual variabilities inthe ecosystem C fluxes. This was coincided with air temperature in the early growingseason, the soil temperature and the vegetation cover. Elevated CO_2significantly decreasedNEE and evapotranspiration (ET), but enhanced GEP, ER and water use efficiency (WUE).Elevated CO_2reduced NEE because the stimulation caused by the elevated CO_2had agreater impact on ER than on GEP. The addition of N stimulated ecosystem C fluxes inboth years and ameliorated the negative impact of elevated CO_2on NEE. Future elevatedCO_2may favor carbon sequestration in C.angusitifoila wetland when coupled withincreasing nitrogen deposition.
(4) Soil respiration displayed strong seasonal patterns, with higher values observed inmidsummer and lower values in spring and autumn in all treatments, which mainly due tohigher atmospheric temperature induced higher plant growth and soil microbial activity inthe midsummer. There were significantly exponential relationships between soil respirationand soil temperature. Elevated CO_2significantly enhanced soil respiration by repeatedmeasures ANOVA. The effect of the N addition on soil respiration was significantly increased in the first year, whereas was decreased in the second year when analyzing yearby year. It was indicated that the effect of the N addition on soil respiration had beeninhibited over time. There was a significantly interactive effect between elevated CO_2andN addition on soil respiration, and the combined effect of them significantly increased soilrespiration by20%-29%compared to that in the control chambers.The results suggest thatelevated CO_2will accelerate soil respiration and carbon cycling in C.angusitifoila wetlandregardless of nitrogen deposition level.
（1）CO_2（550ppm和700ppm）升高条件下，小叶章出现光合下调现象，施氮减缓了光合下调，高氮处理下小叶章无明显的光合适应现象。在实验设计的氮沉降范围内，CO_2浓度升高降低了小叶章叶片气孔导度（Gs）、蒸腾速率（Tr），促进了水分利用效率（WUE）的提高。CO_2浓度升高显著影响了小叶章最大净光合速率（Amax）、光补偿点（LCP）、光饱和点（LSP）、最大羧化速率（Vc,max）和CO_2补偿点（г*）。光合参数对CO_2浓度升高的响应依赖于供氮水平，未施氮和低氮条件下，CO_2升高降低了小叶章叶片Amax、Vc,max，提高了LCP、LSP和г*；在高氮条件下，CO_2浓度升高未显著影响光合参数Amax、暗呼吸速率（Rd）、表观量子产量（AQE）、LCP、LSP、Vc,max和最大电子速率（Jmax）。施氮显著增加了Amax和AQE，降低了Rd、LCP和LSP，促进了小叶章光合能力的提高。CO_2浓度升高显著影响了小叶章叶片光合色素含量。未施氮和低氮条件下，CO_2浓度升高降低了小叶章叶片叶绿素a、叶绿素b、类胡萝卜素和叶绿素总量；但在高氮条件下，小叶章光合色素含量则未发生显著变化。施氮处理显著增加了小叶章叶片光合色素含量。上述结果表明，充足的氮供应可减轻甚至抑制小叶章对CO_2浓度升高的适应。小叶章出现光合适应与Vc,max的降低有关，暗示小叶章发生光合适应很大程度上是由于1,5-二磷酸核酮糖（RuBP）羧化限制。
（2）CO_2浓度升高显著增加了小叶章根中淀粉含量、各器官可溶性糖含量，但对小叶章叶片、茎的淀粉含量的影响不显著。CO_2浓度升高对小叶章叶片可溶性蛋白含量依据氮处理而异，在未施氮条件下，CO_2升高降低了可溶性蛋白含量；在低氮和高氮水平下，则增加了可溶性蛋白含量。施氮处理显著增加了小叶章茎、根中淀粉含量和总淀粉含量、小叶章根、茎和器官总可溶性糖含量以及小叶章叶片可溶性蛋白含量；对叶片中淀粉和可溶性糖含量影响不显著。
不同种类矿质元素对CO_2浓度升高的响应不同。CO_2浓度升高未对小叶章叶片C含量产生显著影响；对小叶章叶片N含量影响因氮处理不同而异。在未施氮和低氮处理下，CO_2升高降低了小叶章叶片N素含量，在高氮处理条件下，小叶章叶片N含量变化不大。CO_2浓度升高显著增加了小叶章叶片P和K含量；显著改变了小叶章C/N、C/P、N/P和N/K。施氮处理显著增加了小叶章叶片N含量；但不同程度降低了小叶章叶片P和K含量，增加了小叶章N/P和N/K。表明CO_2浓度升高和施氮改变了植物养分格局，暗示未来CO_2升高将有利于湿地植物减轻P、K元素限制。
（3）在2010-2011两个生长季中，小叶章湿地净生态系统CO_2交换量（NEE）、生态系统呼吸量（ER）和总生态系统生产力（GEP）呈现夏季高而春秋两季低的季节动态，而且有显著的年际动态，这与生长季早期大气温度、土壤温度和植被盖度有关。CO_2浓度升高降低了NEE、生态系统水分蒸发蒸腾损失量（ET），但增加了ER、GEP和生态系统水分利用效率（WUE）。NEE的降低主要是由于CO_2升高导致ER增加的幅度高于GEP增加的幅度。施氮不仅增加了生态系统碳通量，而且也削弱了CO_2浓度升高对NEE的负面效应。我们的研究结果表明，在未来伴随氮沉降逐渐增加的趋势下，CO_2浓度升高将有利于三江平原小叶章湿地碳固定。
（4）土壤呼吸的季节动态呈现夏季高的单峰曲线模式，主要归因于夏季较高的大气温度促进了植物生长和土壤微生物活性。本研究中，土壤呼吸速率和土壤温度呈现指数相关。CO_2浓度升高条件下，土壤呼吸呈现增加的趋势。施氮处理第一年土壤呼吸显著增加，而第二年土壤呼吸呈现降低的趋势，这表明施氮对土壤呼吸的促进作用随着时间的推移减弱甚至产生抑制作用。CO_2浓度升高和氮沉降对土壤呼吸的影响有显著的互作效应，二者互作显著增加了土壤呼吸，增加幅度在20%-29%。表明在未来CO_2浓度升高条件下，不管氮沉降水平如何，都将增加三江平原小叶章湿地土壤呼吸和土壤碳循环。
The increasingly increases of atmospheric CO_2and nitrogen deposition can influenceplant photosynthetic capacity and then affect ecosystem structure and functions. Thelong-term continuous simulation of atmospheric CO_2and nitrogen (N) depositionexperiments is significant to predict the future global climate change on wetland ecosystemproductivity and carbon sequestration. This study is based on the results through thecontrolled situ-experiment on the Sanjiang Plain wetland in northeast China. This studyfocus on the photosynthetic characteristics, photosynthetic product and mineral elements ofthe dominant species Calamagrostis angustifolia, and ecosystem carbon, water fluxes andsoil respiration of C.angustifolia wetland under different CO_2and N level, which provide atheoretical basis for the wetland adaptation process to global change. There were three CO_2concentration accomplished using Open Top Chambers (370ppm AC,550ppm EC1and700ppmEC2), and three N deposition level (0g N/m~2.a, NN;4g N/m~2.a, LN;8g N/m~2.a, HN) suppliedas NH4NO3. Each treatment had three replicates. The main conclusions are as follows:
(l) Photosynthetic acclimation phenomena occured when C.angustifolia exposed toelevated CO_2(550ppm and700ppm). Nitrogen addition weakened photosyntheticacclimation phenomena and photosynthetic acclimation phenomena were not found underhigh nitrogen addition. The stomatal conductance (Gs) and transpiration rate (Tr) weredecreased and the water use efficiency (WUE) was enhanced under elevated CO_2. ElevatedCO_2significantly influnced the maximum net photosynthetic rate (Amax), lightcompensation point (LCP), light saturation point (LSP), maximum carboxylation rate(Vc,max) and CO_2compensation point (г*). The response of photosynthetic parameters toelevated CO_2varied according to the level of N. Elevated CO_2decreased Amaxand Vc,max,increased LCP, LSP and г*under NN and LN treatment. However, elevated CO_2could notsignificantly affect Amax, dark respiration rate (Rd), apparent quantum efficiency (AQE),LCP, LSP, Vc,maxand maximum electron transport rate (Jmax) under HN. At the same time,N-addition enhanced Amaxand AQE, but decreased Rd,LCP and LSP, which showed that Naddition could increased photosynthetic capacity.
Elevated CO_2significantly altered the photosynthetic pigment content of leaves inC.angustifolia. The response of photosynthetic pigment content to elevated CO_2variedaccording to the level of N. The chlorophyll a, chlorophyll b, carotenoids and totalchlorophyll content of C.angustifolia were decreased under NN and LN treatment.However, the photosynthetic pigments contents of C.angustifolia grown under elevatedCO_2were not significantly lower than those grown under ambient CO_2under HN treatment.N addition enhanced chlorophyll a, chlorophyll b, carotenoids and total chlorophyll contentof C.angustifolia.
The results indicate that the photosynthetic capacity of C.angustifolia declined underlong-term elevated CO_2, but enough N addition could alleviate photosynthetic acclimation.The photosynthetic acclimation occured accompanying with significant decrease of Vc,max.Our results imply that photosynthetic acclimation under elevated CO_2is largely due toRuBP carboxylation limitation.
(2) Elevated CO_2significantly increased the starch content of root, the soluble sugarcontent of leaves, stem, root and total organ in C.angusitifoila, but had no significanteffects on the starch content of leaves and stem. The response of soluble protein content toelevated CO_2varied according to the level of nitrogen. Elevated CO_2decreased solubleprotein content of leaves in C.angusitifoila under NN treatment, but increased solubleprotein content under LN and HN treatment. N addition enhanced the starch content andsoluble sugar content of root, stem and total organ, and the soluble protein content ofleaves, but had no significant effect on the starch content and soluble sugar content ofleaves in C.angusitifoila.
The response of the mineral element to elevated CO_2varied according to the elementsspecies and the level of N. Elevated CO_2had no significant effects on the organic carbon(C) of leaves in C.angusitifoila. The response of N content of leaves to elevated CO_2variedaccording to the level of nitrogen. Elevated CO_2significantly decreased N content ofleaves in C.angusitifoila under NN and LN level but not under HN treatment. Phosphorus(P) and potassium (K) content of leaves was significantly increased under elevated CO_2.Elevated CO_2significantly influnced C/N, C/P, N/P and N/K of leaves in C.angusitifoila. Naddition significantly increased the content of N, decreased the content of P and K, andincreased N/P and N/K of leaves in C.angusitifoila. Our results show that elevated CO_2andnitrogen addition could change the nutrition pattern of plant and imply that elevated CO_2could facilitate wetland plant to mitigate P and K limitation.
(3) The seasonal dynamics of ecosystem carbon fluxes (NEE, ER and GEP) inC.angusitifoila wetland were high in mid-summer and low in the early and late growingseasons in2010and2011. In addition, there were remarkable interannual variabilities inthe ecosystem C fluxes. This was coincided with air temperature in the early growingseason, the soil temperature and the vegetation cover. Elevated CO_2significantly decreasedNEE and evapotranspiration (ET), but enhanced GEP, ER and water use efficiency (WUE).Elevated CO_2reduced NEE because the stimulation caused by the elevated CO_2had agreater impact on ER than on GEP. The addition of N stimulated ecosystem C fluxes inboth years and ameliorated the negative impact of elevated CO_2on NEE. Future elevatedCO_2may favor carbon sequestration in C.angusitifoila wetland when coupled withincreasing nitrogen deposition.
(4) Soil respiration displayed strong seasonal patterns, with higher values observed inmidsummer and lower values in spring and autumn in all treatments, which mainly due tohigher atmospheric temperature induced higher plant growth and soil microbial activity inthe midsummer. There were significantly exponential relationships between soil respirationand soil temperature. Elevated CO_2significantly enhanced soil respiration by repeatedmeasures ANOVA. The effect of the N addition on soil respiration was significantly increased in the first year, whereas was decreased in the second year when analyzing yearby year. It was indicated that the effect of the N addition on soil respiration had beeninhibited over time. There was a significantly interactive effect between elevated CO_2andN addition on soil respiration, and the combined effect of them significantly increased soilrespiration by20%-29%compared to that in the control chambers.The results suggest thatelevated CO_2will accelerate soil respiration and carbon cycling in C.angusitifoila wetlandregardless of nitrogen deposition level.
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