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.

牦牛粪便对青藏高原东部高寒草甸土壤养分系统的影响

川西北高寒草甸位于青藏高原东部地区，是我国四大牧区之一，也是长江和黄河等江河源区的重要水源涵养功能区。近几十年来，大量牦牛粪便被牧民作为生物能源、肥料或者食用菌产业的原料而利用，为草原生态系统的养分平衡增加了生态风险。鉴于在青藏高原地区针对牦牛粪便的相关研究尚未见报道，本文从粪便和土壤养分相互变化趋势的角度出发，研究了粪便在夏秋两季的分解状况和粪便其下及其周围土壤的养分变化。通过此研究，将有助于了解牦牛粪便在自然状态下的分解速率以及粪便对土壤养分及土壤微生物生物量的影响，为粪便对草地土壤生态系统的贡献提供概念性认识，同时也为高寒草甸草地这种脆弱生态系统的可持续管理提供理论依据。 针对牦牛粪便自身养分释放及其对土壤在时间和空间尺度上的影响，本文通过模拟牛粪堆积，在不同时间和固定区域内对牛粪和土壤进行了夏秋两季的采样测定，分析了牛粪及土壤NO3--N、NH4+-N、速效K、无机P、有机C、全N 和全P 含量随时间的变化趋势。得到如下结果： 1. 在研究区域内，牛粪对草地生态系统具有较强的养分（N、P）贡献能力。据初步统计，其估值大致为氮素699~932 kg ha-1，P 素为110~147 kg ha-1。牛粪（湿重、干重）在夏秋季节的分解速率具有较大差异，夏季显著快于秋季。夏季牛粪湿重、干重在2 个月左右之后分别降为初期的14%和24%，而秋季同期分别降为35%和52%。牛粪养分（NO3--N、NH4+-N、速效K、无机P、有机C、全N 和全P）的分解速率在夏季也要显著快于秋季。秋季经过2 个月左右的分解之后，牛粪以上含量分别降至初始态的32%、60%、36%、64%、58%、63%和43%，远高于夏季的同期水平。 2. 在不同季节，牛粪周围的土壤养分（NO3--N、NH4+-N、速效K 和无机P）含量变化随时间改变呈现相同的趋势。但是，牛粪周围不同远近下土壤养分随时间的动态变化幅度略有不同。粪下土壤养分含量随时间变化波动较大，距粪便越远，随时间变化的变幅越为平缓。总体来说，在夏季由于粪便分解较快，对土壤养分的持续作用时间不及秋季，秋季粪便分解变慢，表现出养分的缓释特征。其次，牛粪对粪下土壤影响的持续时间也长于对周围土壤的作用时间。 3. 粪便对土壤养分（NO3--N、NH4+-N、速效K、无机P）影响的范围在不同季节具有差异。夏季要高于秋季，但对周围土壤养分影响的持续时间低于秋季。在夏季短期内，牛粪对土壤NO3--N 和速效K 含量的影响范围能够超过30cm，而对NH4+-N 和无机P 的影响范围则介于10cm 和30cm 之间。在2 个月左右之后，牛粪对周围土壤养分的影响能力基本消失。在秋季，牛粪对周围土壤养分影响范围难以达到30cm 处。粪便在夏季对其下土壤和周围土壤的有机C、全N 和全P 含量并无显著影响，但在秋季能显著增加其下土壤有机C、全N 和全P 含量。 4. 牛粪在秋季对土壤SMB-C、SMB-N 和SMB-P 含量的影响能够持续2 个月以上，由于秋冬季节牛粪分解缓慢，因此推断这种效应持续时间至少能够1 年左右。另一方面，牛粪在秋季对土壤SMB-C、SMB-N 和SMB-P 含量的影响范围主要集中在其下土壤，而周围的影响效应并不明显。 The grassland on the eastern fringe of Qinghai-Tibetan Plateau was one of the four greatest pasture zones in our country and the main water conservation function zones in the hesastream of Yangtse River and Huanghe river. Rencent years, lots of dung in this area was used as biological energy, fertilizer or material of fungoid growing, leading to high risk of nutrient banlance in grassland ecosystem. In view of the researches on the impact of yak dung in this area are relatively rare, the present study focused on the relationship of dung and soil nutrient transformation in sunmer and autumn, which could profoundly illuminate the mechanism of dung decomposition and the effect of dung on soil chemical properties and soil microbe biomass. The present study also contributed to a basic understand and provided scientific management in the high-frigid ecosystem. Decomposition of yak dung and its effect on soil chemical properties in eastern grassland of Qinghai-Tibetan Plateau were determined. The study simulated the real dung pats, took dung and soil samples at different time and fixed-point in summer and autumn. The samples were analysed for NO3--N, NH4+-N, available K, inorganic P, total organic C (TOC), total N (TN), total P (TP). It was concluded that: 1. In study area, the yak dung supplied to ecosystem substantial nutrient. It is estimated that the N contribution of dung was approximately 699~932 kg ha-1, P contribution was approximately 110~147 kg ha-1. The rate of yak decomposition was more rapid in summer than autumn, the wet and dry weight of yak decreased to 14% and 24% respectively after 2 months when dung excreted in summer, with 35% and 52% in autumn. The content of NO3--N、NH4+-N、available K、inorganic P、TOC、TN and TP in dung decomposed more rapid in summer too. After 2 months when dung excreted in autumn, the content of above nutrient decreased to 32%、60%、36%、64%、58%、63% and 43% respectively，which were significantly higher than summer. 2. The content of NO3--N、NH4+-N、available K and inorganic P in soil around dung had the same transformation trend in each season, whereas it was distinguishing at different gradient of distance from dung, the nutrient in soil below dung had the most significant change while the more far from dung, the less change in soil. It was concluded that the yak dung had prolong impacts on soil in autumn compared with summer, besides, it aslo showed that the yak dung had protract effect on soil below dung compared with soil around dung. 3. The yak dung had expansive impact on soil around dung in summer whereas had relatively short effect compared with autumn. In short-term at summer, there was a significant increase about the content of NO3--N and available K around 30cm radius from dung pat while the content of NH4+-N and inorganic P between 10cm to 30cm. After 2 months, the impact almost disappeared. In autumn, the effect was hard to reach 30cm. The yak dung had no significant effects on the content of TOC、TN and TP in soil below or around dung in summer whereas there was a obvious increase in soil below dung pat in autumn. 4. The duration of effect of yak dung on soil microbial biomass(SMB) C、N and P was at least 2 months, maybe even more than 1 year. On the other hand, the impact of dung on SMB-C、SMB-N and SMB-P mainly acted on soil below dung while no obious effect on soil around dung.

Algorithm Studies on How to Obtain a Conditional Nonlinear Optimal Perturbation (CNOP)

The conditional nonlinear optimal perturbation (CNOP), which is a nonlinear generalization of the linear singular vector (LSV), is applied in important problems of atmospheric and oceanic sciences, including ENSO predictability, targeted observations, and ensemble forecast. In this study, we investigate the computational cost of obtaining the CNOP by several methods. Differences and similarities, in terms of the computational error and cost in obtaining the CNOP, are compared among the sequential quadratic programming (SQP) algorithm, the limited memory Broyden-Fletcher-Goldfarb-Shanno (L-BFGS) algorithm, and the spectral projected gradients (SPG2) algorithm. A theoretical grassland ecosystem model and the classical Lorenz model are used as examples. Numerical results demonstrate that the computational error is acceptable with all three algorithms. The computational cost to obtain the CNOP is reduced by using the SQP algorithm. The experimental results also reveal that the L-BFGS algorithm is the most effective algorithm among the three optimization algorithms for obtaining the CNOP. The numerical results suggest a new approach and algorithm for obtaining the CNOP for a large-scale optimization problem.

Long-term grazing alters species composition and biomass of a shrub meadow on the Qinghai-Tibet Plateau

Livestock grazing has long been the most widespread land use on the Qinghai-Tibet Plateau, one of the world's highest ecosystems. However, there has been increasing concern during recent decades because of the rapid increase in livestock numbers. To assess the possible influences of grazing on the vast grassland, a long-term grazing experiment in a shrub meadow on the northern Qinghai-Tibet Plateau was carried out. The experiment included five treatments with different stocking rates and one non-grazing (N) treatment. After 17 years of grazing, treatment differences were clear. The species composition differed markedly between grazing intensities, with a decrease in palatable grass species and an increase in unpalatable forbs at higher grazing intensities. The species richness and species diversity, however, were not significantly different between treatments. Vegetation height decreased significantly at higher grazing intensities. Total above,ground biomass declined considerably and the biomass of forbs increased significantly under the higher grazing intensities. The amount of litter was significantly lower under the higher grazing intensities. The results suggest that long-term grazing alters the species composition, vegetation height and biomass production of the alpine grassland ecosystem without significantly changing species richness.

Effects of temperature on CO2 exchange between the atmosphere and an alpine meadow

The alpine meadow ecosystem on the Qinghai-Tibetan Plateau is characterized by low temperatures because of its high elevation. The low-temperature environment may limit both ecosystem photosynthetic CO2 uptake and ecosystem respiration, and thus affect the net ecosystem CO2 exchange (NEE). We clarified the low-temperature constraint on photosynthesis and respiration in an alpine meadow ecosystem on the northern edge of the plateau using flux measurements obtained by the eddy covariance technique in two growing seasons. When we compared NEE during the two periods, during which the leaf area index and other environmental parameters were similar but the mean temperature differed, we found that NEE from 9 August to 10 September 2001, when the average temperature was low, was greater than that during the same period in 2002, when the average temperature was high, but the ecosystem gross primary production was similar during the two periods. Further analysis showed that ecosystem respiration was significantly higher in 2002 than in 2001 during the study period, as estimated from the relationship between temperature and nighttime ecosystem respiration. The results suggest that low temperature controlled the NEE mainly through its influence on ecosystem respiration. The annual NEE, estimated from 15 January 2002 to 14 January 2003, was about 290 g CO2 m(-2) year(-1). The optimum temperature for ecosystem NEE under light-saturated conditions was estimated to be around 15 degrees C.

Characterizing evapotranspiration over a meadow ecosystem on the Qinghai-Tibetan Plateau

To characterize evapotranspiration (ET) over grasslands on the Qinghai-Tibetan Plateau, we examined ET and its relevant environmental variables in a Kobresia meadow from 2002 to 2004 using the eddy covariance method. The annual precipitation changed greatly, with 554, 706, and 666 mm a(-1) for the three consecutive calendar years. The annual ET varied correspondingly to the annual precipitation with 341, 407, and 426 mm a(-1). The annual ET was, however, constant at about 60% of the annual precipitation. About 85% annual ET occurred during the growing season from May to September, and the averaged ET for this period was 1.90, 2.23, and 2.22 mm/d, respectively for the three consecutive years. The averaged ET was, however, very low (< 0.40 mm/d) during the nongrowing season from October to April. The annual canopy conductance (gc) and the Priestley-Taylor coefficient (a) showed the lowest values in the year with the lowest precipitation. This study first demonstrates that the alpine meadow ecosystem is characterized by a low ratio of annual ET to precipitation and that the interannual variation of ET is determined by annual precipitation.

Depression of net ecosystem CO2 exchange in semi-arid Leymus chinensis steppe and alpine shrub

Uptake and release of carbon in grassland ecosystems is very critical to the global carbon balance and carbon storage. In this study, the dynamics of net ecosystem CO2 exchange (FNEE) of two grassland ecosystems were observed continuously using the eddy covariance technique during the growing season of 2003. One is the alpine shrub on the Tibet Plateau, and the other is the sem-arid Leymus chinensis steppe in Inner Mongolia of China. It was found that the FNEE of both ecosystems was significantly depressed under high solar radiation. Comprehensive analysis indicates that the depression of FNEE in the L. chinensis steppe was the results of decreased plant photosynthesis and increased ecosystem respiration (R-eco) under high temperature. Soil water stress in addition to the high atmospheric demand under the strong radiation was the primary factor limiting the stomatal conductance. In contrast, the depression of FNEE in the alpine shrub was closely related to the effects of temperature on both photosynthesis and ecosystem respiration, coupled with the reduction of plant photosynthesis due to partial stomatal closure under high temperature at mid-day. The R,c of the alpine shrub was sensitive to soil temperature during high turbulence (u* > 0.2 m s(-1)) but its FNEE decreased markedly when the temperature was higher than the optimal value of about 12 degrees C. Such low optimal temperature contrasted the optimal value (about 20 degrees C) for the steppe, and was likely due to the acclimation of most alpine plants to the long-term low temperature on the Tibet Plateau. We inferred that water stress was the primary factor causing depression of the FNEE in the semi-arid steppe ecosystem, while relative high temperature under strong solar radiation was the main reason for the decrease of FNEE in the alpine shrub. This study implies that different grassland ecosystems may respond differently to climate change in the future. (c) 2006 Elsevier B.V All rights reserved.

Stability of alpine meadow ecosystem on the Qinghai-Tibetan Plateau

The meadow ecosystem on the Qinghai-Tibetan Plateau is considered to be sensitive to climate change. An understanding of the alpine meadow ecosystem is therefore important for predicting the response of ecosystems to climate change. In this study, we use the coefficients of variation (Cv) and stability (E) obtained from the Haibei Alpine Meadow Ecosystem Research Station to characterize the ecosystem stability. The results suggest that the net primary production of the alpine meadow ecosystem was more stable (Cv = 13.18%) than annual precipitation (Cv = 16.55%) and annual mean air temperature (Cv= 28.82%). The net primary production was insensitive to either the precipitation (E = 0.0782) or air temperature (E = 0.1113). In summary, the alpine meadow ecosystem on the Qinghai-Tibetan Plateau is much stable. Comparison of alpine meadow ecosystem stability with other five natural grassland ecosystems in Israel and southern African indicates that the alpine meadow ecosystem on the Qinghai-Tibetan Plateau is the most stable ecosystem. The alpine meadow ecosystem with relatively simple structure has high stability, which indicates that community stability is not only correlated with biodiversity and community complicity but also with environmental stability. An average oscillation cycles of 3-4 years existed in annual precipitation, annual mean air temperature, net primary production and the population size of consumers at the Haibei natural ecosystem. The high stability of the alpine meadow ecosystem may be resulting also from the adaptation of the ecosystem to the alpine environment.

Energy exchange between the atmosphere and a meadow ecosystem on the Qinghai-Tibetan Plateau

To reveal the potential contribution of grassland ecosystems to climate change, we examined the energy exchange over an alpine Kobresia meadow on the northeastern Qinghai-Tibetan Plateau. The annual pattern of energy exchange showed a clear distinction between periods of frozen soil with the daily mean soil temperature at 5 cm (T-s5 &LE; 0 &DEG; C) and non-frozen soil (T-s5 > 0 &DEG; C). More than 80% of net radiation was converted to sensible heat (H) during the frozen soil period, but H varied considerably with the change in vegetation during the non-frozen soil period. Three different sub-periods were further distinguished for the later period: (1) the pre-growth period with Bowen ratio (β) > 1 was characterized by a high β of 3.0 in average and the rapid increase of net radiation associated with the increases of H, latent heat (LE) and soil heat; (2) during the Growth period when β &LE; 1, the LE was high but H fluxes was low with β changing between 0.3 and 0.4; (3) the post-growth period with average β of 3.6 when H increased again and reached a second maximum around early October. The seasonal pattern suggests that the phenology of the vegetation and the soil water content were the major factors affecting the energy partitioning in the alpine meadow ecosystem. © 2005 Elsevier B.V. All rights reserved.

Carbon dioxide exchange between the atmosphere and an alpine meadow ecosystem on the Qinghai-Tibetan Plateau, China

We used the eddy covariance method to measure the M exchange between the atmosphere and an alpine meadow ecosystem (37degrees29-45'N, 101degrees12-23'E, 3250m a.s.l.) on the Qinghai-Tibetan Plateau, China in the 2001 and 2002 growing seasons. The maximum rates Of CO2 uptake and release derived from the diurnal course Of CO2 flux (FCO2) were -10.8 and 4.4 mumol m(-2) s(-1), respectively, indicating a relatively high net carbon sequestration potential as compared to subalpine coniferous forest at similar elevation and latitude. The largest daily CO2 uptake was 3.9 g cm(-2) per day on 7 July 2002, which is less than half of those reported for lowland grassland and forest at similar latitudes. The daily CO2 uptake during the measurement period indicated that the alpine ecosystem might behave as a sink of atmospheric M during the growing season if the carbon lost due to grazing is not significant. The daytime CO2 uptake was linearly correlated with the daily photosynthetic photon flux density each month. The nighttime averaged F-CO2 showed a positive exponential correlation with the soil temperature, but apparently negative correlation with the soil water content. (C) 2004 Elsevier B.V. All rights reserved.