环境CO2浓度和群落密度对红桦(Betula albosinensis Burk.)幼苗碳氮吸收和分配的影响

当前大气CO2浓度升高是全球变化的主要趋势之一，CO2浓度升高还会引起全球变暖等其它环境问题，因而CO2浓度浓度升高对植物影响的研究已经成为全球变化领域的焦点。红桦是川西亚高山地区暗针叶林演替初期的先锋树种和演替后期的建群种，在群落演替过程中它对环境因子的响应决定红桦群落的演替进程。本文通过控制CO2浓度的气候室试验，研究了CO2浓度倍增环境下，不同密度水平红桦碳氮固定、分配可能发生的改变，并探讨了升高大气CO2浓度对群体内部竞争的影响。以期通过本研究明确川西亚高山地区代表性物种红桦对未来气候变化的响应，为今后采取措施应对气候变化、妥善进行森林管理提供理论依据和科学指导。主要研究结果如下： 1.升高CO2浓度对红桦幼苗生长的影响以及树皮、树干响应的不同 (1) CO2浓度升高显著促进红桦幼苗的生物量、株高、基茎的生长，同时也改变生物量在体内的分配格局，主要是增加根和主茎、减少叶在总生物量中的比重。(2)树皮和树干对升高CO2浓度的影响有差异，它们对CO2浓度升高的反应程度不同，但反应方向一致。 2.密度的副效应 (1) 增加种植密度对单株生物量、株高和基径的生长具有副效应，也降低升高CO2浓度对红桦生长的正效应。(2) 增加种植密度，显著增加红桦幼苗的群体生物量，从而使红桦群体固定更多的大气CO2气体。可见密度在决定红桦生物量及固碳能力方面具有重要意义。探索适合未来大气CO2浓度升高条件下植物生长的密度，对未来的森林经济生产、生态恢复具有重要意义。 3. 升高CO2浓度对红桦幼苗苗冠结构及冠层内部竞争的影响 (1) 冠幅、冠高、苗冠表面积和苗冠体积等树冠特征均受CO2浓度升高的影响而增加，但是受密度增加的影响而降低。(2) 单位苗冠投影面积叶片数(LDcpa)和单位苗冠体积叶片数(LDcv)均低于相应的现行CO2浓度处理，这主要是由于冠幅和冠高的快速生长所造成的。(3) LDcpa和LDcv的降低表明，红桦在升高CO2浓度的条件下，会作出积极的响应，从而缓解由于群体和个体生长的增加所引起的竞争压力的增加。 4. 升高CO2浓度对红桦幼苗养分元素吸收与分配的影响 (1) CO2浓度升高，植株各器官N、P含量降低，但单株N、P总吸收量均增加。红桦幼苗体内N、P浓度的下降是由于生物量迅速增加引起的稀释效应造成的。(2) CO2浓度升高，N、P向主茎和根的分配增加，向叶片的分配减少，主要是由于前者在总生物量中的比重增加，而后者减少了。(3) CO2浓度升高，氮磷利用效率(NUE和PUE)提高，氮磷累积速率(NAcR和PAcR)显著增加。而NUE和PUE的提高可以有效缓解CO2浓度升高后，亚高山和高山地区森林土壤中养分元素不足对森林生产力的限制。 5. 升高CO2浓度对红桦幼苗群体碳平衡的影响 (1) 升高CO2浓度对植物的光合作用、呼吸速率和生长均具有促进作用。(2) 土壤有机碳含量在实验前期迅速增加，后期积累速率下降。(3) 升高CO2浓度以后，土壤呼吸显著增强；土壤呼吸还具有明显的季节变化。(4) 红桦群体日固碳量受到升高CO2浓度的促进作用。结果(1)-(4)说明所研究群落的碳动态对现行的气候波动是敏感的；所研究群落在作为大气CO2气体的源-汇关系方面至少存在季节间的源汇飘移。(5)种植密度的升高显著增加了群体固碳量。 6. 升高CO2浓度对红桦幼苗生长后期叶片衰老的影响 升高CO2浓度有利于减缓红桦幼苗叶片生长季节末期的衰老。生长季节末期，随着CO2浓度的升高光合速率和可溶性蛋白含量均呈上升趋势，同时MDA（丙二醛）含量下降，保护酶SOD（超氧化物岐化酶）、CAT（过氧化氢酶）活性升高。由此说明，升高CO2浓度有利于减缓生长季节后期叶片的衰老，使叶片维持较高的光合速率，也从生理学的角度支持了本文及前人有关CO2浓度升高促进植物光合和生长的假说及结果。 The increased CO2 concentration is one of the most important problems among global changes. The increase of CO2 will also cause other environmental problems, such as global warming, etc. So the effects of elevated CO2 on plant have drawn sights of many scientists in the research field of global change. Red birch (Betula albosinensis) usually emerges as the pioneer species in initial stage and as constructive species in later stages of forest community succession of the dark coniferous forests in Western Sichuan, China. It’s response to elevated CO2 may determine the succession process of the community where it lives in. By controlling CO2 at the ambient and twice as the ambient level (ambient + 350 umol mol-1) using enclosed-top chambers (ETC), possible effects of elevated CO2 on carbon fixation and allocation under two plantation densities are investigated. The effects of elevated CO2 on competition within canopy of red birch seedlings are also observed in the present paper. We hope to make sure of the effects of elevated CO2 on the representative species, red birch. And so that, our results could provide a strong theoretical evidence and scientific direction for forest management and afforestation under a future, CO2 elevated world. The results are as fowllows: 1. The effects of elevated CO2 on growth and the different responses of wood and bark of red birch seedlings (1) Elevated CO2 increases the growth of seedling biomass, seedling height and basal diameter of red birch. It also changed the biomass allocation in red birch seedlings. The ratio of root and main stem to all biomass is increased and the ratio of leaf is decreased. (2) Tree bark and wood show different response degree but similar response direction to elevated CO2. 2. Negative effects of planting density (1) The increase of planting density showes negative effects on the individual growth of seedling biomass, seedling height and basal diameter of red birch. It also eliminates the positive effects of elevated CO2 on growth of red birch seedlings. (2) Community biomass is increased by the elevated planting density, which means that the high density red birch community could fix more CO2 than the low density one. These results show that planting density plays an important role in determining biomass and carbon fixation ability of red birch community. Thus, exploring proper planting density becomes economically important for the future, CO2 elevated word. 3. The effects of elevated CO2 on crown architecture and competition within canopy of red birch seedlings (1) Crown width, crown depth, crown surface area and crown volume are all increased under the influence of elevated CO2. (2) Leaf number per unit area of projected crown area (LDcpa) and per unit volume of crown volume (LDcv) are lower under elevated CO2. This is resulted from the stimulated growth of tree crown features. (3) The decrease of LDcpa and LDcv indicate that plants will respond forwardly to reduce the possible increase of competition resulted from stimulated growth of individual plant and collectives in conditions of elevated CO2. 4. The effects of elevated CO2 on nutrition accumulation and allocation of red birch seedlings (1) Contents of N and P decrease due to the prompt increase of biomass of plant organs caused by elevated CO2. However, their accumulations increase under elevated CO2. (2) Elevated CO2 increases the allocation of N, P to main stem but reduced its allocation to leaf for that dry weight of the former increased but the dry weight of the later decreased. (3) Using efficiencies of N, P (NUE and PUE) and their accumulation rates (NAcR and PAcR) are found to increase under elevated CO2. Soil nutrition contents are always the limiting factors for plant growth at subalpine and alpine region. The increased NUE and PUE are helpful to eliminate the nutrition limitation in this area in the future world, when CO2 concentration doubles the ambient. 5. The effects of elevated CO2 on carbon balance of red birch communities (1) Net photosynthetic rates (Pn), dark respiration rates (Rd) and growth are all stimulated by elevated CO2. (2) Content soil organic carbon increases sharply at the primary stage of experiments and then the increasing rates decrease to a low level at later stages. (3) Soil respiration rates increase significantly with the elevation of CO2 concentration. (4) The daily carbon fixations of whole community are heightened by elevated CO2. The results (1)-(4) suggest that, the community being studied are sensitive to current climate change; the studied community, as a sink of atmospheric CO2, is pool-sink alternative between seasons. (5) The carbon fixations are increased along the increase of planting densities. 6. The effects of elevated CO2 on physiological features of leaf senescences of red birch seedlings at the later stage of growing season Elevated CO2 helps to postpone the leaf senescences of red birch at the end of the growth season. CO2 enrichment increases the photosynthetic rates, contents of soluble proteins and photosynthetic pigments. And meanwhile contents of malondialdehyde (MDA) decreases and activities of superoxide dismutase (SOD) and catalase (CAT) are both increased. These results suggest that the senescences of red birch leaves are delayed by elevated CO2, which keep the photosynthetic rates at relatively high levels. Our results lend supports to hypothesis and results on stimulated photosynthetic rates and growth from both other researchers and the present paper.

模拟增温对川西亚高山人工云杉林土壤碳含量和土壤呼吸的影响

全球气候变化已经成为不争的事实，其中全球变暖是近年来国内外的研究热点之一。土壤碳库作为陆地生态系统最大的碳库，气温升高必然会导致一系列的土壤碳储量和碳通量的变化，这些微小的变化又可能导致大气CO2浓度的变化并强化这种变暖的趋势。目前，土壤碳循环对温度升高的响应仍然是陆地碳循环研究最缺乏的部分，对土壤有机碳动态变化的研究仍存在着很大的不确定性与争议。四川西部的亚高山人工针叶林是青藏高原东部高寒林区的重要组成部分，是研究全球变化对森林生态系统影响的关键地区和重要森林类型。本研究通过采用原位人工模拟增温装置（Open-top chambers，OTCs）对川西米亚罗60年人工云杉林土壤实施增温，研究高海拔地区森林，尤其是人工森林系统下的土壤有机碳 含量、土壤呼吸及土壤酶活性对温度升高的响应。结果表明： 1. 增温处理的660天（2005年11月至2007年9月）期间，增温条件下的平均气温和土壤平均温度分别比对照提高0.43 ℃和0.27 ℃；0~10 cm土壤含水量在增温的不同时期均有不同程度的降低。 2. 土壤蔗糖酶、蛋白酶和脲酶活性在温度升高的不同阶段均有不同程度的提高。在增温处理300天（2006.09）、540天（2007.05）、600天（2007.07）和660天（2007.05）后，0~10 cm层的蔗糖酶活性分别比对照提高了36.36%（P<0.05）、24.31%、14.54%（P<0.05）和7.22%，脲酶活性分别提高了12.90%、24.19%（P<0.01）、34.48%（P<0.05）和14.64%(P<0.05)，蛋白酶活性分别提高了31.37%、1.99%、3.70%和17.80%。10~20 cm层的土壤酶活性也均有不同程度的提高，但均没有显著差异。蔗糖酶、脲酶和蛋白酶活性均呈现出随土层加深而减弱的趋势。 3. 土壤过氧化氢酶和多酚氧化酶活性在增温的第1年内均有不同程度的提高，但在增温的第2年内比对照有所降低。增温300天后（2006.09），过氧化氢酶和多酚氧化酶在0~10 cm层分别比对照增加3.76%和49.25%（P<0.05），10~20 cm层分别增加了5.54%和29.67%。在增温的第2年内，增温540天（2007.05）、600天（2007.07）和660天（2007.09）后，0~10 cm层的过氧化氢酶活性分别比对照降低了27.70%（P<0.05）、4.34%和1.47%，多酚氧化酶活性分别降低了5.86%、11.76%（P<0.05）和7.47%。增温的第2年内，10~20 cm层的过氧化氢酶和多酚氧化酶活性也均有不同程度的降低，但差异均未达到显著水平。不同土层之间相比较，过氧化氢酶活性随土层加深而降低，多酚氧化酶活性随土层加深而增加。 4. 土壤有机碳和有机质在增温的不同阶段，含量比对照均有所降低；且随增温时间的延长，降低的幅度下降。0~10 cm层的土壤有机碳和土壤有机质在增温300天（2006.09）、540天（2007.05）、600天（2007.07）和660天（2007.09）后分别降低了8.69%、4.35%、3.80%和2.44%，差异均未达到显著水平。土壤全氮含量在增温后与对照相比无明显的增加或者降低趋势。增温条件下的土壤C/N比与对照相比有所降低，但在增温各阶段的差异均不显著。10~20 cm层的有机碳、有机质和C/N比也有不同程度的降低趋势，但差异均不显著。不同土层之间相比，0~10 cm层的有机碳、有机质、全氮含量和C/N比均高于10~20 cm层，呈现出随土层加深而降低的趋势。 5. 土壤呼吸速率在增温第1年内，与对照相比明显提高，但在增温处理2年后，与对照相比无显著变化。增温300天（2006.09）和360天（2006.11）后分别提高了13.32%和21.17%，差异显著。增温处理540天（2007.05）到660天（2007.09）期间，与对照相比，不仅没有明显的提升，反而有些月份比对照有所降低，对温度升高的敏感性降低，呈现出对温度升高的适应性。土壤呼吸的日呼吸速率呈现单峰曲线形式，在14:00~20:00期间达到最大值，在4:00~10:00期间具有最低值。土壤呼吸的季节变化，呈现出与外界环境温度相一致的趋势，在7月份（夏季） 最高，11月份（冬季）最低。土壤呼吸与2 cm土壤温度、5 cm土壤温度和空气温度均呈极显著指数相关，与0~10 cm土壤含水量呈线性相关，相关性达到显著水平，但低于土壤呼吸与温度的相关性。 The past century has seen a marked increase in atmospheric carbon dioxide concentrations and a concomitant warming that has drawn scientific attention to the link between global carbon stocks and climate change. In particular, the decomposition and turnover of soil organic matter is recognised as an important determinant of carbon driven climate change. The slightly variation in soil organic carbon will result in the increase of atmospheric carbon dioxide concentrations and reinforce the tendency of warming. The experiment was conducted in Subalpine coniferous forest in western Sichuan province. Subalpine coniferous forest in western Sichuan was a important part of eastern Qinghai-Tibetan Plateau, which play a important role in reseaching the sensitivity of forest ecosystem to climate change. To investigate the effects of elevated temperature on soil organic carbon content, soil respiration rates, and soil enzyme activities in subalpine Picea asperata plantations, a esimulated warming measure was applied with Open-top chambers. The results were as followed: 1) During the period from Nov. 2005 to Sep. 2007, mean air temperature and soil temperature were respectively 0.43℃ and 0.27℃ the ambient higher. Soil water content decreased to different exent in different months in warmed plots than in unwarned plots at depth of 0-10 cm. 2) In general, elevated temperature enhanced the soil enzyme activities of invertase, protease, and urease. In the first year of warming—after 300 days’ treatment (in Sep,2006), the activities of invertase, protease, and urease increased by 36.36%, 12.90% and 31.37% respectively at the depths of 0-10 cm，among which the activity of invertase reached statistic significance. In the second year of warming, invertase activity increased by 24.31% after 540 days’ treament (in May, 2007), 14.54% after 600 days’ treament (in Jul, 2007) and 7.22% after 660 days’ treatment (in Sep, 2007) at the depths of 0-10 cm, and the differences in July and Septemmber were statistically significant. Elveated temperature also increased the activity of urease in the second year of warming and had significant effects in May and July. The activity of protease in warmed plots was also higher than in unwarmed plots at depths of 0-10 cm, but there was no significant difference. Elevated temperature had no significant effects on all soil enzyme acitivities at the depths of 10-20 cm in the first and sencond year. The values of above-mentioned soil enzyme all decreased with soil layers. 3) Eleavted temperature enhanced the activities of catalase and polyphenol oxidase in the first year of warming while they turned out downtrend in the second year. The activity of catalase increased by 3.76% and 5.54% at depths of 0-10 cm and 10-20 cm respectively in the first year—after 300 days’ warming (in Sep, 2006), the differences of which had no statistical significance. The activity of polyphenol oxidase was significantly increased by 49.25% at depths of 0-10 cm and not significantly increased by 29.67% at depths of 10-20 cm after 300 days’ warming. In the second year of warming, the catalase activity was significantly decreased by 27.70% after 540 days’ treament (in May, 2007) and not significantly decreased by 4.34% and 1.47% after 600 days’ (in Jul, 2007) and 660 days’ treament (in Sep, 2007) respectively. The activities of catalase and polyphenol oxidase at depths of 10-20 cm were decreased to different extent, but there was no significant difference. Catalase activity stepped down with soil layers while polyphenol oxidase activity stepped up. 4) Increased temperature in both the first year and the second year resulted tendency of decrease in the contents of soil organic carbon and soil organic matter, and C/N ratios at soil depths of 0-10 cm and 10-20 cm. However, with the prolonged warming, the tendency of decrease gradually tapered off and the extent of decrease in the second year of experiment were lower than that in the first year. The contents of soil organic carbon and soil organic matter were all decreased 8.69% by warming in the first year and dcreased 4.35%, 3.80% and 2.44% in May, July and September of the second year, but no significant difference were found. The C/N ratios increased 8.52% in the first year of warming and had less increment in the second year, all of which were not statistical significant. Eleveated temperature had no obvious effect on the content of tatol N in two year consecutive warming experiment. The contents of soil organic carbon and soil organic matter, total N and C/N ratios all had the tendency of dcreasing with soil layers. 5) Soil respiration rates were significantly enhanced by 13.32% and 21.17% after 300 days’ (in Sep, 2006) and 360 days’ (in Nov, 2006) treament in the first year of warming, but the same showed no obvious difference in the second year of treatment, which was assumed the adaptability of soil respiration with a certain heightened temperature. Diurnal soil resspiration showed a daily variation with a minimum value between 4:00 and 10:00 h and a maximum value between 14:00 and 20:00 h, coinciding with the minimum and maximum values of soil temperature at 2 cm. Soil respiration rates exhibited a pronounced seasonal variation with minimum values in Novmber and a maximum value in July, approximately coinciding with the seasonal variation of air and soil temperature. An exponential function provided the best fit for soil respiration with temperature while a quadric equation was used to estimate the effect of soil moisture on soil respiration, which were all significantly correlated. Soil respiraion rate was more highly correlated with the soil temperature than soil moisture.

川西北高寒草甸土壤呼吸研究：季节动态和放牧效应

除植被冠层的光合作用之外，土壤的呼吸作用是陆地生态系统碳收支中最大的通量。土壤呼吸即使发生较小的变化也能显著地减缓或加剧大气中CO2浓度的增加，从而明显影响到全球气候变化。土壤呼吸速率变化与否以及变化的方向可以反映生态系统对环境变化的敏感程度和响应模式。尽管如此，土壤呼吸仍是一个为人们了解不多的生态系统过程。 草地生态系统是陆地生态系统的一个重要组成部分。针对草地土壤呼吸进行野外实验研究和相应方法论的探讨将对区域乃至全球碳源汇性质的准确估算具有重要的科学意义。然而，近几年来关于草地土壤呼吸的主要研究工作都集中在温带草原和部分热带草原，而针对高寒草甸生态系统土壤呼吸的研究报道还很少。 2008年4月至2009年4月期间，我分别在2008年6、8、10、12月和2009年2月和4月分6次对川西北的典型高寒草甸群落的土壤呼吸进行观测，分析了不同类型高寒草甸群落土壤呼吸的季节变化特征以及环境因子和放牧模式对其影响。主要研究结果如下： 1）该地区高寒草甸生态系统在生长季（6月～8月）土壤呼吸速率较大（6.07～9.30μmolCO2¡m-2¡s-1 ） ， 在非生长季（ 12 月～ 2 月） 较小（ 0.16 ～0.49μmolCO2¡m-2¡s-1 ） 。土壤CO2 年累积最大释放量为3963 ～ 5730gCO2¡m-2¡yr-1，其中，生长季土壤CO2的释放量占年总释放量的85%～90%。非生长季占10%～15%。非生长季所占比例略小于冬季积雪覆盖地区的冬季土壤呼吸占年土壤呼吸量的比例（14%～30%）。温度，尤其地温，是影响该地区高寒草甸生态系统土壤呼吸速率的最主要环境因子。土壤呼吸速率与地上生物量和土壤水分之间没有显著相关性，但是土壤含水量过大会导致土壤呼吸速率下降。 2）在观测期内，草丘区的土壤呼吸显著高于对照区的土壤呼吸，其最大土壤呼吸速率为16.77μmolCO2¡m-2¡s-1，土壤CO2 年累积最大释放量为8145gCO2¡m-2¡yr-1，是对照区的近2 倍。由于草丘在高寒草甸中占有较大的面积比例（近30%），因此，它将对高寒草甸生态系统的碳循环起着重要的作用。 3）放牧模式不仅可以影响高寒草甸群落的土壤CO2 排放，而且还可以改变土壤呼吸的温度敏感性（Q10）。本研究表明，在生长季有长期放牧活动干扰时将会增加土壤向大气中释放二氧化碳的速度，促使土壤碳库中碳的流失。禁牧样地的土壤呼吸速率在刚禁牧时先迅速增大，随着禁牧时间的延长土壤呼吸速率将会下降。此外，与其它放牧模式相比，冬季放牧将高寒草甸群落土壤呼吸速率在生长季达到最大值的时间明显向后推迟。不同放牧模式下高寒草甸群落土壤呼吸的Q10 值大小顺序为：禁牧一年群落>冬季放牧群落>禁牧三年群落>夏季放牧群落>自由放牧群落。 4）基于呼吸室技术的观测方法中，测量前的剪草处理可以明显改变该地区高寒草甸群落的土壤温度和土壤呼吸速率。在生长季，剪草处理将使土壤呼吸速率的瞬时响应增加90%左右。由于剪草处理明显增加了剪草样方白天的土壤温度，而土壤温度与土壤呼吸之间存在着极显著的指数相关关系，因而剪草处理导致土壤呼吸速率迅速增加。因此，在高寒地区基于呼吸室技术观测的土壤呼吸应当进行校正。 综上所述，川西北高寒草甸生态系统土壤呼吸速率在生长季较高，而在非生长季较低。土壤温度是影响该地区土壤呼吸的最主要环境因子。在实验观测期，草丘区土壤呼吸速率显著高于对照区的，是对照区土壤呼吸速率的近2倍。由于测量前的剪草处理可以明显改变待测点的土壤呼吸速率，因此，应对在高寒地区基于呼吸室技术观测的土壤呼吸进行校正。 Soil respiration is the second largest component (less than plant phtotosynthesis) of carbon dioxide flux between terrestrial ecosystems and the atmosphere. A minor change in soil respiration rate can significantly slow down or accelerate the increase of atmospheric CO2 concentration that is closely related to global climatic change. In turn, the change in the flux direction and rate of soil respiration may indicate the elasticity and stability of ecosystems to global changes and human disturbance. However, soil respiration is still an ecosystem process that has been poorly understood. Grassland ecosystem is an important component of the terrestrial ecosystem. Accurately estimating the CO2 flux from soil to atmosphere in situ is the key to evaluating the carbon resource and sink regionally or globally. Despite of extensive studies on the temperate and tropic grasslands, the soil respiration of alpine meadows has not substantially been measured. In the current study, soil respiration was measured for an annual cycle from April, 2008 to April, 2009 for the alpine meadow in northwestern Sichuan Province of China to determine the seasonal variation of soil respiration for the typical plant communities. The results are shown as follows: 1) Large seasonal variation of soil respiration was observed in the alpine meadow. The rate of soil respiration was the greatest (6.07～9.30μmolCO2¡m-2¡s-1) in June and the smallest (0.16 ～ 0.49μmolCO2¡m-2¡s-1) occurred from December to February in the non-growing season. The total emission of soil CO2 was estimated as 3963～5730 gCO2¡m-2¡yr-1, 85%~90% of which was released during the growing season, and 10%~15% during the non-growing season which was slightly less than the ratio of winter and annual CO2 flux from soil. Temperature, particularly the soil temperature, was the major environmental factor regulating the soil respiration. Significant and positive relationships were not found between soil respiration and soil moisture and between soil respiration and plant above-ground biomass, but excessive soil water content would decrease in the rate of soil respiration. 2) The rate of soil respiration in grass hummock communities was up to 16.77μmolCO2¡m-2¡s-1, which was about twice as great as in the controls (communities located in low and even sites). Considering the large proportion (about 30% on average) of hummock area in the meadow, it can be concluded that the hummocks played an important role in the carbon cycling of the study ecosystem. 3) Grazing patterns affected the flux of CO2 emission and the temperature sensitivity of soil respiration (Q10) in the alpine meadow. Grazing during growing season increased the rate of soil respiration. The rate of soil respiration increased significantly immediately after the alpine meadow being fenced, but thereafter decreased. In addition, grazing in winter delayed the peak respiration rate relative to the non-grazing mode. The Q10 value was the largest in the non-grazed area for one year, and next came the area with grazing in winter, followed by the non-grazed area for three years, the area with grazing in summer, and the non-limited grazed area. 4) In the chamber-based techniques, clipping manipulation before each measurement increased the transient rate of soil respiration by about 90% in the summer of the alpine meadow. As increase in soil temperature at daytime in the clipped plots by clipping and the exponential relationship between soil respiration and temperature, clipping manipulation led to increase in the rate of soil respiration. This suggested that a correction should be done for the techniques if employed in alpine and cold regions. In summary, the rate of soil respiration in the alpine meadow was the greatest in June and the smallest occurred from ecember to February in the non-growing season. Soil temperature was the major environmental factor regulating the soil respiration. The rate of soil respiration in grass hummock communities was up to 16.77μmolCO2¡m-2¡s-1, which was about twice as great as in the controls. A correction should be done for the techniques if employed in alpine and cold regions, because of the effect of clipping manipulation on soil temperature and respiration.

连作对设施黄瓜产量和品质及土壤酶活性的影响

揭示黄瓜产生连作障碍的机理,为设施蔬菜的可持续发展和减轻连作障碍提供参考依据。【方法】在日光温室内进行田间试验,以露地玉米地土壤为对照,研究不同连作年限(4,5,8和12年)下黄瓜产量和品质及土壤酶活性的变化。【结果】随着黄瓜连作年限的增加,黄瓜产量、可溶性固形物和维生素C含量均下降,硝酸盐含量上升,土壤呼吸强度降低,土壤脲酶活性、蔗糖酶活性、碱性磷酸酶活性均呈先上升后降低的趋势;随着黄瓜生长季节的变化,土壤脲酶活性呈先上升后降低再升高的趋势,土壤蔗糖酶活性、碱性磷酸酶活性和土壤呼吸强度均呈先上升后降低的趋势。【结论】随着连作年限的延长,设施黄瓜的产量和品质均下降,土壤脲酶、蔗糖酶和碱性磷酸酶活性均呈先升高后降低的趋势。

Environmental controls over carbon exchange of three forest ecosystems in eastern China

National Natural Science Foundation of China [30590381, 30670384]; Knowledge Innovation Project of the Chinese Academy of Sciences [KZCX2-YW-432]; National Key Research and Development Program [2002CB412501]; 'Hundred Talents' Program of the Chinese Acade

Comparative study on CO2 emissions from different types of alpine meadows during grass exuberance period

Potentilla fruticosa scrub, Kobresia humilis meadow and Kobresia tibetica meadow are widely distributed on the Qinghai-Tibet Plateau. During the grass exuberance period from 3 July to 4September, based on close chamber-GC method, a study on CO2 emissions from different treatments was conducted in these meadows at Haibei research station, CAS. Results indicated that mean CO2emission rates from various treatments were 672.09+152.37 mgm-2h-1 for FC (grass treatment); 425.41+191.99 mgrn-2h-1 for FJ (grass exclusion treatment); 280.36+174.83 mgrn-2h-1 for FL (grass and roots exclusion treatment); 838.95+237.02 mgm-2h-1 for GG (scrub+grass treatment); 528.48+205.67 mgm-2h-1for GC (grass treatment); 268.97 ±99.72 mgm-2h-1 for GL (grass and roots exclusion treatment); and 659.20±94.83 mgm-2h-1 for LC (grass treatment), respectively (FC, FJ, FL, GG, GC, GL, LC were the Chinese abbreviation for various treatments). Furthermore, Kobresia humilis meadow, Potentilla fruticosa scrub meadow and Kobresia tibetica meadow differed greatly in average CO2 emission rate of soil-plant system, in the order of GG＞FC＞LC＞GC. Moreover, in Kobresia humilis meadow,heterotrophic and autotrophic respiration accounted for 42% and 58% of the total respiration of soil-plant system respectively, whereas, in Potentilla fruticosa scrub meadow, heterotrophic and autotrophic respiration accounted for 32% and 68% of total system respiration from G-G; 49% and 51%from GC. In addition, root respiration from Kobresia humilis meadow approximated 145 mgCO2m-2h-1,contributed 34% to soil respiration. During the experiment period, Kobresia humilis meadow and Potentilla fruticosa scrub meadow had a net carbon fixation of 111.11 grn-2 and 243.89 grn-2,respectively. Results also showed that soil temperature was the main factor which influenced CO2 emission from alpine meadow ecosystem, significant correlations were found between soil temperature at 5 cm depth and CO2 emission from GG, GC, FC and FJ treatments. In addition, soil moisture may be the inhibitory factor of CO2 emission from Kobresia tibetica meadow, and more detailed analyses should be done in further research.

CO2H2O and energy exchange of an Inner Mongolia steppe ecosystem during a dry and wet year

We used an eddy covariance technique to measure evapotranspiration and carbon flux over two very different growing seasons for a typical steppe on the Inner Mongolia Plateau, China. The rainfall during the 2004 growing season (344.7 mm) was close to the annual average (350.43 mm). In contrast, precipitation during the 2005 growing season was significantly lower than average (only 126 mm). The wet 2004 growing season had a higher peak evapotranspiration (4 mm day(-1)) than did the dry 2005 growing season (3.3 mm day(-1)). In 2004, latent heat flux was mainly a consumption resource for net radiation, accounting for similar to 46% of net radiation. However, sensible heat flux dominated the energy budget over the whole growing season in 2005, accounting for 60% of net radiation. The evaporative rate (LE/R-n) dropped by a factor of four from the non-soil stress to soil water limiting conditions. Maximum half-hourly CO2 uptake was -0.68 mg m(-2) s(-1) and maximum ecosystem exchange was 4.3 g CO2 m(-2) day(-1) in 2004. The 2005 drought growing stage had a maximum CO2 exchange value of only -0.22 mg m(-2) s(-1) and a continuous positive integrated-daily CO2 flux over the entire growing season, i.e. the ecosystem became a net carbon source. Soil respiration was temperature dependent when the soil was under non-limiting soil moisture conditions, but this response declined with soil water stress. Water availability and a high vapor pressure deficit severely limited carbon fixing of this ecosystem; thus, during the growing season, the capacity to fix CO2 was closely related to both timing and frequency of rainfall events. (c) 2007 Published by Elsevier Masson SAS.

西藏高原草原化嵩草草甸生态系统呼吸及碳平衡

The soil respiration and net ecosystem productivity of Kobresia littledalei meadow ecosystem was investigated at Dangxiong grassland station, one grassland field station of Lhasa Plateau Ecosystem Research Station. Soil respiration and soil heterotrophic respiration were measured at the same time by using Li6400-09 chamber in growing season of year 2004. The response of soil respiration and its components, i.e. microbial heterotrophic respiration and root respiration to biotic and abiotic factors were addressed. We studied the daily and seasonal variation on Net Ecosystem carbon Exchange (NEE) measured by eddy covariance equipments and then the regression models between the NEE and the soil temperature. Based on the researches, we analyzed the seasonal variation in grass biomass and estimated NEE combined the Net Ecosystem Productivity with heterogeneous respiration and then assessed the whether the area is carbon source or carbon sink. 1.Above-ground biomass was accumulated since the grass growth started from May; On early September the biomass reached maximum and then decreased. The aboveground net primary production (ANPP) was 150.88 g m~" in 2004. The under-ground biomass reached maximum when the aboveground start to die back. Over 80% of the grass root distributed at the soil depth from 0 to 20cm. The underground NPP was 1235.04 g m"2.. Therefore annual NPP wasl.385X103kg ha"1, i.e.6236.6 kg C ha"1. 2. The daily variation of soil respiration showed single peak curve with maximum mostly at noon and minimum 4:00-6:00 am. Daily variations were greater in June, July and August than those in September and October. Soil respiration had strong correlation with soil temperature at 5cm depth while had weaker correlation with soil moisture, air temperature, surface soil temperature, and so on. But since early September the soil respiration had a obviously correlation with soil moisture at 5cm depth. Biomass had a obviously linearity correlation with soil respiration at 30th June, 20th August, and the daytime of 27th September except at 23lh October and at nighttime of 27th September. We established the soil respiration responding to the soil temperature and to estimate the respiration variation during monsoon season (from June through August) and dry season (May, September and October). The regression between soil respiration and 5cm soil temperature were: monsoon season (June through August), Y=0.592expfl()932\ By estimating , the soil daily respiration in monsoon season is 7.798gCO2m"2 and total soil respiration is 717.44 gCC^m" , and the value of Cho is 2.54; dry season (May, September and October), Y=0.34exp°'085\ the soil daily respiration is 3.355gCO2m~2 and total soil respiration is 308.61 gCC^m", and the value of Cho is 2.34. So the total soil respiration in the grown season (From May to October) is 1026.1 g CO2IT1"2. 3. Soil heterogeneous respiration had a strong correlation with soil temperature especially with soil temperature at 5cm depth. The variation range in soil heterogeneous respiration was widely. The regression between soil heterogeneous respiration and 5cm soil temperature is: monsoon season, Y=0.106exp ' 3x; dry season, Y=0.18exp°"0833x.By estimating total soil heterotrophic respiration in monsoon season is 219.6 gCC^m"2, and the value of Cho is 3.78; While total soil heterogeneous respiration in dry season is 286.2 gCCbm"2, and the value of Cho is 2.3. The total soil heterotrophic respiration of the year is 1379.4kg C ha"1. 4. We estimated the root respiration through the balance between soil respiration and the soil heterotrophic respiration. The contribution of root respiration to total respiration was different during different period: re-greening period 48%; growing period 69%; die-back period 48%. 5. The Ecosystem respiration was relatively strong from May to October, and of which the proportion in total was 97.4%.The total respiration of Ecosystem was 369.6 g CO2 m" .we got the model of grass respiration respond to the soil temperature at 5cm depth and then estimated the daytime grass respiration, plus the nighttime NEE and daytime soil respiration. But when we estimated the grass respiration, we found the result was negative, so the estimating value in this way was not close. 6. The estimating of carbon pool or carbon sink. The NPP minus the soil heterogeneous respiration was the NEE, and it was 4857.3kg C o ha"1, which indicated that the area was the carbon sink.

Carbon distribution within the plant and rhizosphere in laboratory and field-grown Lolium perenne at different stages of development

Carbon distribution within perennial ryegrass was determined at different stages of plant development, by pulse-labelling laboratory and field-grown plants with ¹⁴C-CO₂. During the early stages of growth (23-51 days), C distribution of laboratory grown plants was not markedly affected by plant age, with 12.4-24% of net assimilated label lost into the soil as root-soil respiration. The percentage of net assimilate translocated below ground was 20-28% during this stage of growth. At 65 days, the percentage of the label translocated below ground decreased to 8.1% of the net assimilate, with a subsequent decrease in root-soil respiration to 3.9%. The ability of the plant to fix the label (expressed in MBq g^-1 oven dry total plant weight) decreased steadily as the plants aged. When the 30 day old plants were subjected to water stress (soil water potential -1.5 MPa) for 2 days before pulse-labelling, root-soil respiration of the pulse-label decreased compared with plants grown at field capacity. The distribution of a ¹⁴C pulse-label within perennial ryegrass grown under field conditions was found to be dependent on the age of the plants. For 4 week old plants, 67% of net assimilated label was translocated below ground, with 64.8% of this respired by the roots and soil. Less label was translocated below ground at subsequent pulse-labels from weeks 8 to 24. The proportion of label translocated below ground respired by the roots and soil also decreased. The investment of label in the plant shoots was found to be greater in field grown plants as compared to plants of the same age grown in a controlled, laboratory environment. © 1990.

Distribution of assimilated carbon within the plant and rhizosphere of Lolium perenne:Influence of temperature

Perennial rye-grass plants were pulse labelled with [¹⁴C]-CO₂ over a range of temperatures (5-25°C). The fate of the label was determined within the plant and soil. The temperature at which plants were pulse labelled had a marked effect on the distribution of the label within the plant and soil system. Root-soil respiration increased from 5.7 to 24.15% when expressed as a percentage of net assimilated label. The percentage of label remaining in the plant root and in the soil was greater at 5 and 25°C, with a minimum for both these components at 15°C. At 15°C the percentage of net assimilated label that remained in the shoots was greater than at other temperatures, with this percentage decreasing at the lower and higher temperatures. © 1989.

Wirkung von Ernterückständen transgener Pflanzen auf die mikrobielle C- und N-Transformation in landwirtschaftlich genutzten Böden am Beispiel von Bt-Mais

In der vorliegenden Arbeit ging es um die Erarbeitung, Anwendung und Beurteilung von quantitativen Analysenverfahren / Methoden für ein Monitoring von durch Bt-Mais verursachbaren Umwelteffekten im Boden. Die Ausgangsthese besagte, dass sich transgene Maisstreu beim mikrobiellen Abbau anders verhält als konventionelle. Bezugnehmend auf die These wurden zwei Freilandversuche (Freilandmikrokosmenmethode nach Raubuch 1997 über 2 Jahre, Quantifizierung des Maisstreuabbaus mit Hilfe kleiner Bodensäulen über 1 Jahr) und zwei Inkubationsversuche im Labor (INK bei drei verschiedenen Temperaturen über 49 Tage und INK mit verschiedenen landwirtschaftlich genutzten Böden über 49 Tage mit jeweils kontinuierlicher Respirationsratenermittlung nach Isermeyer 1952) sowie Inhaltsstoffbestimmungen der Maisstreu durchgeführt. Für alle Untersuchungen wurde Streu der vier Maissorten Novelis (transgen, Monsanto 810), Nobilis (Isolinie von Novelis), Valmont (transgen, Bt 176, Fa. Syngenta) und Prelude (Isolinie von Valmont) eingesetzt. Nach Beendigung der Laborversuche sowie des Freilandversuches nach der Freilandmikrokosmenmethode wurden mikrobielle Messgrößen wie Adenylategehalt, Ergosterolgehalt, Cmik- und Nmik-Gehalt am Boden-Streu-Gemisch bestimmt. Der Einsatz der Isotopentechnik (Bestimmung von 13C/12C an gemahlenem Boden-Streu-Gemisch bzw. gefriergetrocknetem K2SO4 als Extrakt aus dem Boden-Streu-Gemisch) ermöglichte eine genaue Quantifizierung der abgebauten Maisstreu und brachte dadurch Aufschluss über das Abbauverhalten verschiedener Maissorten. Bezüglich der Ermittlung der mikrobiellen Messgrößen ergab sich für die transgene Sorte Novelis* stets eine durchschnittlich geringere pilzliche Biomasse. Langfristig ergaben sich bei der Kohlenstoff- und Stickstoffdynamik keine Trends hinsichtlich transgener bzw. konventioneller Maisstreu. Sowohl im Freilandversuch nach der Mikrokosmenmethode als auch in den Inkubationsversuchen trat das Phänomen der kurzzeitigen Respirationsratenerhöhung der Mikroorganismen nach Zugabe der transgenen Maissorten auf, welches nicht bei Zugabe der konventionellen Maisstreu auszumachen war. ______________________________

Field measurements of the CO2 evolution rate under different crops during an irrigation cycle in a mountain oasis of Oman

For millennia oasis agriculture has been the backbone of rural livelihood in the desertic Sultanate of Oman. However, little is known about the functioning of these oasis systems, in particular with respect to the C turnover. The objective was to determine the effects of crop, i.e. alfalfa, wheat and bare fallow on the CO2 evolution rate during an irrigation cycle in relation to changes in soil water content and soil temperature. The gravimetric soil water content decreased from initially 24% to approximately 16% within 7 days after irrigation. The mean CO2 evolution rates increased significantly in the order fallow (27.4 mg C m^−2 h^−1) < wheat (45.5 mg C m^−2 h^−1) < alfalfa (97.5 mg C m^−2 h^−1). It can be calculated from these data that the CO2 evolution rate of the alfalfa root system was nearly four times higher than the corresponding rate in the wheat root system. The decline in CO2 evolution rate, especially during the first 4 days after irrigation, was significantly related to the decline in the gravimetric water content, with r = 0.70. CO2 evolution rate and soil temperature at 5 cm depth were negatively correlated (r = -0.56,n = 261) due to increasing soil temperature with decreasing gravimetric water content.

Application of biogas slurries from energy crops to arable soils and their impact on carbon and nitrogen dynamics

The use of renewable primary products as co-substrate or single substrate for biogas production has increased consistently over the last few years. Maize silage is the preferential energy crop used for fermentation due to its high methane (CH4) yield per hectare. Equally, the by-product, namely biogas slurry (BS), is used with increasing frequency as organic fertilizer to return nutrients to the soil and to maintain or increase the organic matter stocks and soil fertility. Studies concerning the application of energy crop-derived BS on the carbon (C) and nitrogen (N) mineralization dynamics are scarce. Thus, this thesis focused on the following objectives: I) The determination of the effects caused by rainfall patterns on the C and N dynamics from two contrasting organic fertilizers, namely BS from maize silage and composted cattle manure (CM), by monitoring emissions of nitrous oxide (N2O), carbon dioxide (CO2) and CH4 as well as leaching losses of C and N. II) The investigation of the impact of differences in soil moisture content after the application of BS and temperature on gaseous emissions (CO2, N2O and CH4) and leaching of C and N compounds. III) A comparison of BS properties obtained from biogas plants with different substrate inputs and operating parameters and their effect on C and N dynamics after application to differently textured soils with varying application rates and water contents. For the objectives I) and II) two experiments (experiment I and II) using undisturbed soil cores of a Haplic Luvisol were carried out. Objective III) was studied on a third experiment (experiment III) with disturbed soil samples. During experiment I three rainfall patterns were implemented including constant irrigation, continuous irrigation with periodic heavy rainfall events, and partial drying with rewetting periods. Biogas slurry and CM were applied at a rate of 100 kg N ha-1. During experiment II constant irrigation and an irrigation pattern with partial drying with rewetting periods were carried out at 13.5°C and 23.5°C. The application of BS took place either directly before a rewetting period or one week after the rewetting period stopped. Experiment III included two soils of different texture which were mixed with ten BS’s originating from ten different biogas plants. Treatments included low, medium and high BS-N application rates and water contents ranging from 50% to 100% of water holding capacity (WHC). Experiment I and II showed that after the application of BS cumulative N2O emissions were 4 times (162 mg N2O-N m-2) higher compared to the application of CM caused by a higher content of mineral N (Nmin) in the form of ammonium (NH4+) in the BS. The cumulative emissions of CO2, however, were on the same level for both fertilizers indicating similar amounts of readily available C after composting and fermentation of organic material. Leaching losses occurred predominantly in the mineral form of nitrate (NO3-) and were higher in BS amended soils (9 mg NO3--N m-2) compared to CM amended soils (5 mg NO3--N m-2). The rainfall pattern in experiment I and II merely affected the temporal production of C and N emissions resulting in reduced CO2 and enhanced N2O emissions during stronger irrigation events, but showed no effect on the cumulative emissions. Overall, a significant increase of CH4 consumption under inconstant irrigation was found. The time of fertilization had no effect on the overall C and N dynamics. Increasing temperature from 13.5°C to 23.5°C enhanced the CO2 and N2O emissions by a factor of 1.7 and 3.7, respectively. Due to the increased microbial activity with increasing temperature soil respiration was enhanced. This led to decreasing oxygen (O2) contents which in turn promoted denitrification in soil due to the extension of anaerobic microsites. Leaching losses of NO3- were also significantly affected by increasing temperature whereas the consumption of CH4 was not affected. The third experiment showed that the input materials of biogas plants affected the properties of the resulting BS. In particular the contents of DM and NH4+ were determined by the amount of added plant biomass and excrement-based biomass, respectively. Correlations between BS properties and CO2 or N2O emissions were not detected. Solely the ammonia (NH3) emissions showed a positive correlation with NH4+ content in BS as well as a negative correlation with the total C (Ct) content. The BS-N application rates affected the relative CO2 emissions (% of C supplied with BS) when applied to silty soil as well as the relative N2O emissions (% of N supplied with BS) when applied to sandy soil. The impacts on the C and N dynamics induced by BS application were exceeded by the differences induced by soil texture. Presumably, due to the higher clay content in silty soils, organic matter was stabilized by organo-mineral interactions and NH4+ was adsorbed at the cation exchange sites. Different water contents induced highest CO2 emissions and therefore optimal conditions for microbial activity at 75% of WHC in both soils. Cumulative nitrification was also highest at 75% and 50% of WHC whereas the relative N2O emissions increased with water content and showed higher N2O losses in sandy soils. In summary it can be stated that the findings of the present thesis confirmed the high fertilizer value of BS’s, caused by high concentrations of NH4+ and labile organic compounds such as readily available carbon. These attributes of BS’s are to a great extent independent of the input materials of biogas plants. However, considerably gaseous and leaching losses of N may occur especially at high moisture contents. The emissions of N2O after field application corresponded with those of animal slurries.

Linking carbon and sulphur cycling during simulated drought cycles in peat from six sites across the UK

Water table draw-down is thought to increase peat decomposition and, therefore, DOC release. However, several studies have shown lower DOC concentrations during droughts relative to ‘normal’ periods with high water table. We carried out controlled incubation experiments at 10°C on 10x10 cm peat soil cores collected from six UK sites across a sulphur deposition gradient. Our aim was to quantify the balance between microbial consumption and chemical precipitation of DOC due to episodic acidification driven by sulphur redox reactions by comparing changes in soil water chemistry to microbial activity (i.e. soil respiration and trace gas fluxes). During dry periods, all sites showed a concurrent increase in SO4 and soil respiration and a decline in DOC. However, the magnitude of change in both DOC and SO4 varied considerably between sites according to historical sulphur deposition loads and the variation in acid/base chemistry.

Processes controlling DOC in pore water during simulated drought cycles in six different UK peats

The effect of episodic drought on dissolved organic carbon (DOC) dynamics in peatlands has been the subject of considerable debate, as decomposition and DOC production is thought to increase under aerobic conditions, yet decreased DOC concentrations have been observed during drought periods. Decreased DOC solubility due to drought-induced acidification driven by sulphur (S) redox reactions has been proposed as a causal mechanism; however evidence is based on a limited number of studies carried out at a few sites. To test this hypothesis on a range of different peats, we carried out controlled drought simulation experiments on peat cores collected from six sites across Great Britain. Our data show a concurrent increase in sulphate (SO4) and a decrease in DOC across all sites during simulated water table draw-down, although the magnitude of the relationship between SO4 and DOC differed between sites. Instead, we found a consistent relationship across all sites between DOC decrease and acidification measured by the pore water acid neutralising capacity (ANC). ANC provided a more consistent measure of drought-induced acidification than SO4 alone because it accounts for differences in base cation and acid anions concentrations between sites. Rewetting resulted in rapid DOC increases without a concurrent increase in soil respiration, suggesting DOC changes were primarily controlled by soil acidity not soil biota. These results highlight the need for an integrated analysis of hydrologically driven chemical and biological processes in peatlands to improve our understanding and ability to predict the interaction between atmospheric pollution and changing climatic conditions from plot to regional and global scales.