Determinants of root-associated fungal communities within Asteraceae in a semi-arid grassland

Summary

1.While plant–fungal interactions are important determinants of plant community assembly and ecosystem functioning, the processes underlying fungal community composition are poorly understood.
2.Here, we studied for the first time the root-associated eumycotan communities in a set of co-occurring plant species of varying relatedness in a species-rich, semi-arid grassland in Germany. The study system provides an opportunity to evaluate the importance of host plants and gradients in soil type and landscape structure as drivers of fungal community structure on a relevant spatial scale. We used 454 pyrosequencing of the fungal internal transcribed spacer region to analyse root-associated eumycotan communities of 25 species within the Asteraceae, which were sampled at different locations within a soil type gradient. We partitioned the variance accounted for by three predictors (host plant phylogeny, spatial distribution and soil type) to quantify their relative roles in determining fungal community composition and used null model analyses to determine whether community composition was influenced by biotic interactions among the fungi.
3.We found a high fungal diversity (156 816 sequences clustered in 1100 operational taxonomic units (OTUs)). Most OTUs belonged to the phylum Ascomycota (35.8%); the most abundant phylotype best-matched Phialophora mustea. Basidiomycota were represented by 18.3%, with Sebacina as most abundant genus. The three predictors explained 30% of variation in the community structure of root-associated fungi, with host plant phylogeny being the most important variance component. Null model analysis suggested that many fungal taxa co-occurred less often than expected by chance, which demonstrates spatial segregation and indicates that negative interactions may prevail in the assembly of fungal communities.
4.Synthesis. The results show that the phylogenetic relationship of host plants is the most important predictor of root-associated fungal community assembly, indicating that fungal colonization of host plants might be facilitated by certain plant traits that may be shared among closely related plant species.

Potential soil carbon sequestration in overgrazed grassland ecosystems

Excessive grazing pressure is detrimental to plant productivity and may lead to declines in soil organic matter. Soil organic matter is an important source of plant nutrients and can enhance soil aggregation, limit soil erosion, and can also increase cation exchange and water holding capacities, and is, therefore, a key regulator of grassland ecosystem processes. Changes in grassland management which reverse the process of declining productivity can potentially lead to increased soil C. Thus, rehabilitation of areas degraded by overgrazing can potentially sequester atmospheric C. We compiled data from the literature to evaluate the influence of grazing intensity on soil C. Based on data contained within these studies, we ascertained a positive linear relationship between potential C sequestration and mean annual precipitation which we extrapolated to estimate global C sequestration potential with rehabilitation of overgrazed grassland. The GLASOD and IGBP DISCover data sets were integrated to generate a map of overgrazed grassland area for each of four severity classes on each continent. Our regression model predicted losses of soil C with decreased grazing intensity in drier areas (precipitation less than 333 mm yr(-1)), but substantial sequestration in wetter areas. Most (93%) C sequestration potential occurred in areas with MAP less than 1800 mm. Universal rehabilitation of overgrazed grasslands can sequester approximately 45 Tg C yr(-1), most of which can be achieved simply by cessation of overgrazing and implementation of moderate grazing intensity. Institutional level investments by governments may be required to sequester additional C.

Validating MODIS-derived land surface evapotranspiration with in situ measurements at two AmeriFlux sites in a semiarid region

Reducing uncertainties in the estimation of land surface evapotranspiration (ET) from remote-sensing data is essential to better understand earth-atmosphere interactions. This paper demonstrates the applicability of temperature-vegetation index triangle (T-s-VI) method in estimating regional ET and evaporative fraction (EF, defined as the ratio of latent heat flux to surface available energy) from MODIS/Terra and MODIS/Aqua products in a semiarid region. We have compared the satellite-based estimates of ET and EF with eddy covariance measurements made over 4 years at two semiarid grassland sites: Audubon Ranch (AR) and Kendall Grassland (KG). The lack of closure in the eddy covariance measured surface energy components is shown to be more serious at MODIS/Aqua overpass time than that at MODIS/Terra overpass time for both AR and KG sites. The T-s-VI-derived EF could reproduce in situ EF reasonably well with BIAS and root-mean-square difference (RMSD) of less than 0.07 and 0.13, respectively. Surface net radiation has been shown to be systematically overestimated by as large as about 60 W/m(2). Satisfactory validation results of the T-s-VI-derived sensible and latent heat fluxes have been obtained with RMSD within 54 W/m(2). The simplicity and yet easy use of the T-s-VI triangle method show a great potential in estimating regional ET with highly acceptable accuracy that is of critical significance in better understanding water and energy budgets on the Earth. Nevertheless, more validation work should be carried out over various climatic regions and under other different land use/land cover conditions in the future.

Diurnal, seasonal and annual variation in net ecosystem CO2 exchange of an alpine shrubland on Qinghai-Tibetan plateau

Thus far, grassland ecosystem research has mainly been focused on low-lying grassland areas, whereas research on high-altitude grassland areas, especially on the carbon budget of remote areas like the Qinghai-Tibetan plateau is insufficient. To address this issue, flux of CO2 were measured over an alpine shrubland ecosystem (37 degrees 36'N, 101 degrees 18'E; 325 above sea level [a. s. l.]) on the Qinghai-Tibetan Plateau, China, for 2 years (2003 and 2004) with the eddy covariance method. The vegetation is dominated by formation Potentilla fruticosa L. The soil is Mol-Cryic Cambisols. To interpret the biotic and abiotic factors that modulate CO2 flux over the course of a year we decomposed net ecosystem CO2 exchange (NEE) into its constituent components, and ecosystem respiration (R-eco). Results showed that seasonal trends of annual total biomass and NEE followed closely the change in leaf area index. Integrated NEE were -58.5 and -75.5 g C m(-2), respectively, for the 2003 and 2004 years. Carbon uptake was mainly attributed from June, July, August, and September of the growing season. In July, NEE reached seasonal peaks of similar magnitude (4-5 g C m(-2) day(-1)) each of the 2 years. Also, the integrated night-time NEE reached comparable peak values (1.5-2 g C m(-2) day(-1)) in the 2 years of study. Despite the large difference in time between carbon uptake and release (carbon uptake time < release time), the alpine shrubland was carbon sink. This is probably because the ecosystem respiration at our site was confined significantly by low temperature and small biomass and large day/night temperature difference and usually soil moisture was not limiting factor for carbon uptake. In general, R-eco was an exponential function of soil temperature, but with season-dependent values of Q(10). The temperature-dependent respiration model failed immediately after rain events, when large pulses of R-eco were observed. Thus, for this alpine shrubland in Qinghai-Tibetan plateau, the timing of rain events had more impact than the total amount of precipitation on ecosystem R-eco and NEE.

Carbon flow in an upland grassland: effect of liming on the flux of recently photosynthesized carbon to rhizosphere soil

The effect of liming on the flow of recently photosynthesized carbon to rhizosphere soil was studied using (CO2)-C-13 pulse labelling, in an upland grassland ecosystem in Scotland. The use of C-13 enabled detection, in the field, of the effect of a 4-year liming period of selected soil plots on C allocation from plant biomass to soil, in comparison with unlimed plots. Photosynthetic rates and carbon turnover were higher in plants grown in limed soils than in those from unlimed plots. Higher delta(13)C% values were detected in shoots from limed plants than in those from unlimed plants in samples clipped within 15 days of the end of pulse labelling. Analysis of the aboveground plant production corresponding to the 4-year period of liming indicated that the standing biomass was higher in plots that received lime. Lower delta(13)C% values in limed roots compared with unlimed roots were found, whereas no significant difference was detected between soil samples. Extrapolation of our results indicated that more C has been lost through the soil than has been gained via photosynthetic assimilation because of pasture liming in Scotland during the period 1990-1998. However, the uncertainty associated with such extrapolation based on this single study is high and these estimates are provided only to set our findings in the broader context of national soil carbon emissions.

The origins of C4 grasslands: integrating evolutionary and ecosystem science.

The evolution of grasses using C4 photosynthesis and their sudden rise to ecological dominance 3 to 8 million years ago is among the most dramatic examples of biome assembly in the geological record. A growing body of work suggests that the patterns and drivers of C4 grassland expansion were considerably more complex than originally assumed. Previous research has benefited substantially from dialog between geologists and ecologists, but current research must now integrate fully with phylogenetics. A synthesis of grass evolutionary biology with grassland ecosystem science will further our knowledge of the evolution of traits that promote dominance in grassland systems and will provide a new context in which to evaluate the relative importance of C4 photosynthesis in transforming ecosystems across large regions of Earth.

Nitrogen and carbon dynamics in grassland soils and plants after application of digestate

A better understanding of effects after digestate application on plant community, soil microbial community as well as nutrient and carbon dynamics is crucial for a sustainable grassland management and the prevention of species and functional diversity loss. The specific research objectives of the thesis were: (i) to investigate effects after digestate application on grass species and soil microbial community, especially focussing on nitrogen dynamic in the plant-soil system and to examine the suitability of the digestate from the “integrated generation of solid fuel and biogas from biomass” (IFBB) system as fertilizer (Chapter 3). (ii) to investigate the relationship between plant community and functionality of soil microbial community of extensively managed meadows, taking into account temporal variations during the vegetation period and abiotic soil conditions (Chapter 4). (iii) to investigate the suitability of IFBB-concept implementation as grassland conservation measure for meadows and possible associated effects of IFBB digestate application on plant and soil microbial community as well as soil microbial substrate utilization and catabolic evenness (Chapter 5). Taken together the results indicate that the digestate generated during the IFBB process stands out from digestates of conventional whole crop digestion on the basis of higher nitrogen use efficiency and that it is useful for increasing harvestable biomass and the nitrogen content of the biomass, especially of L. perenne, which is a common species of intensively used grasslands. Further, a medium application rate of IFBB digestate (50% of nitrogen removed with harvested biomass, corresponding to 30 50 kg N ha-1 a-1) may be a possibility for conservation management of different meadows without changing the functional above- and belowground characteristic of the grasslands, thereby offering an ecologically worthwhile alternative to mulching. Overall, the soil microbial biomass and catabolic performance under planted soil was marginally affected by digestate application but rather by soil properties and partly by grassland species and legume occurrence. The investigated extensively managed meadows revealed a high soil catabolic evenness, which was resilient to medium IFBB application rate after a three-year period of application.

Plant diversity effects on grassland productivity are robust to both nutrient enrichment and drought

Global change drivers are rapidly altering resource availability and biodiversity. While there is consensus that greater biodiversity increases the functioning of ecosystems, the extent to which biodiversity buffers ecosystem productivity in response to changes in resource availability remains unclear. We use data from 16 grassland experiments across North America and Europe that manipulated plant species richness and one of two essential resources—soil nutrients or water—to assess the direction and strength of the interaction between plant diversity and resource alteration on above-ground productivity and net biodiversity, complementarity, and selection effects. Despite strong increases in productivity with nutrient addition and decreases in productivity with drought, we found that resource alterations did not alter biodiversity–ecosystem functioning relationships. Our results suggest that these relationships are largely determined by increases in complementarity effects along plant species richness gradients. Although nutrient addition reduced complementarity effects at high diversity, this appears to be due to high biomass in monocultures under nutrient enrichment. Our results indicate that diversity and the complementarity of species are important regulators of grassland ecosystem productivity, regardless of changes in other drivers of ecosystem function.

Antelope mating strategies facilitate invasion of grasslands by a woody weed

Intra and interspecific variation in frugivore behaviour can have important consequences for seed dispersal outcomes. However, most information comes from among-species comparisons, and within-species variation is relatively poorly understood. We examined how large intraspecific differences in the behaviour of a native disperser, blackbuck antelope Antilope cervicapra, influence dispersal of a woody invasive, Prosopis juliflora, in a grassland ecosystem. Blackbuck disperse P. juliflora seeds through their dung. In lekking blackbuck populations, males defend clustered or dispersed mating territories. Territorial male movement is restricted, and within their territories males defecate on dung-piles. In contrast, mixed-sex herds range over large areas and do not create dung-piles. We expected territorial males to shape seed dispersal patterns, and seed deposition and seedling recruitment to be spatially localized. Territorial males had a disproportionately large influence on seed dispersal. Adult males removed twice as much fruit as females, and seed arrival was disproportionately high on territories. Also, because lek-territories are clustered, seed arrival was spatially highly concentrated. Seedling recruitment was also substantially higher on territories compared with random sites, indicating that the local concentration of seeds created by territorial males continued into high local recruitment of seedlings. Territorial male behaviour may, thus, result in a distinct spatial pattern of invasion of grasslands by the woody P. juliflora. An ex situ experiment showed no beneficial effect of dung and a negative effect of light on seed germination. We conclude that large intraspecific behavioural differences within frugivore populations can result in significant variation in their effectiveness as seed dispersers. Mating strategies in a disperser could shape seed dispersal, seedling recruitment and potentially plant distribution patterns. These mating strategies may aid in the spread of invasives, such as P. juliflora, which could, in turn, negatively influence the behaviour and ecology of native dispersers.

内蒙古温带草原生态系统过程对气候变化响应的实验研究

羊草草原、大针茅草原和草甸草原是内蒙古温带典型草原地区的三种主要植被类型，本文以此为研究对象，应用时空替代、温室、网室、自然降水接移、养分添加等多种方法模拟未来可能发生的气候变化，研究了这些变化对上述草原植物群落和生态系统过程的直接和间接影响。主要研究结果如下： 1. 气候变暖及其诱导的土壤养分释放速率增加对植物群落有明显的影响，表现在种种植物群落在高度、盖度、密度和生物量等都有不同程度的变化。说明我国温带草原生态系统对气候变化反应敏感。 2. 在群落说平上，气候变暖直接或间接使地上生物量提高17%～90%，但在种群水平，不同种群对模拟气候变化的响应具有很强的个性，而不具普遍性，不同种群反应的大小、方向不同。因此由在不同群落进行研究得出的推论只有部分是可能的，并且即使观测到相类似的反应，它们内在的机制也很可能是不同的。 3. 气候变化显著地影响着凋落物的分解特征。但不同的气候变化情景下，凋落物分解过程的响应不同。在温度升高降水变化不大或升高的情景下，凋落物的分解速率将加快;在气温上升，降水明显下降的情景下，凋落物的分解速率将降低。 4. 草甸草原土壤碳素释放过程和氮素矿化过程对气候变暖有明显的响应。气候变暖将促进土壤有机碳的释放，使草甸草原土壤成为碳源，同时加速氮素矿化速率，在一定程度上提高土壤有效养分的浓度。 5. 在中小尺度上，海拔样带所是应用时空替代原理，研究陆地生态系统与气候变化的理想平台，生物量是综合反应气候变化对土壤-植物系统影响的敏感指标。 6. 草原土壤有机碳和氮在2m～3m的小尺度上存在空间自相关性，其空间自相关尺度受人类活动的影响。

锡林河流域温带草地植物VOC释放及其对草地生态系统碳循环的贡献

植物源挥发性有机碳化合物（Volitale organic compounds, VOC）是大气VOC的主要来源，与对流层大气质量、大气化学密切相关。鉴于温带草地的分布范围很广，草地植物VOC释放潜力某种程度上影响植物源VOC的总释放量。另外，植物源VOC也是光合作用固定碳素的损失方式之一，可能在特定区域或生态系统中具有重要意义。基于上述想法，本文设计了四个方面的实验作为研究内容：1） 温带草地物种水平VOC释放潜力、及其与植物功能群的关系？2) 沙地植物物种水平VOC释放潜力、及其与植物功能群的关系？3) 沙地植物-草地植物VOC释放潜力存在显著性差异吗？4) 温带典型草地和退化草地的VOC释放速率如何？在生态系统水平，植物源VOC对温带草地碳循环的贡献多大？ 在所测定的175种温带草地植物中，不同植物间异戊二烯和单萜释放潜力差异很大;除少数物种外，大多数植物的异戊二烯和单萜释放潜力都较低，尤其是典型草地的优势物种。在此基础上，作者探讨了分类学赋值方法对温带草地植被的可行性，并初步建立了锡林河流域温带草地植物的VOC释放目录（共277种植物）。另外，温带草地植物的异戊二烯和单萜释放潜力与植物功能群（植物生活型和水分功能群）具有一定的联系，尤其是植物生活型。总的来说，温带草原的优势生活型（物种），即多年生根茎禾草（或多年生丛生禾草），具有较低的异戊二烯和单萜释放潜力。各水分功能群间差异不显著，但中旱生植物、旱中生植物 (温带草原的优势功能群)，具有较低的异戊二烯、单萜释放潜力。因此，温带草原退化过程中，那些具有较高VOC释放潜力的植物，重要性将会增加。 沙地植物种类组成非常丰富，不同物种间的异戊二烯和单萜释放潜力变异也很大。另外，沙地植物的异戊二烯和单萜释放潜力与其功能群间关系较密切，不同植物生活型间差异显著;其中也以多年生根茎禾草、多年生从生禾草的释放潜力最低，而乔木的释放潜力相对最高;该结论基本与草地的研究结论一致。然而，沙地植物的异戊二烯和单萜释放潜力与其水分功能群的关系比较模糊，中生植物具有更高的释放潜力，湿生植物的释放潜力较小。 通过对比沙地植物和草地植物的释放潜力，发现沙地植物的异戊二烯和单萜释放潜力比草地植物高，且这种差异整体上显著。另外，这种显著性差异，在不同植物生活型、水分功能群间也同样存在。沙地植物比对应的草地植物具有更高的异戊二烯和单萜释放潜力，最可能的原因：沙地正午的温度明显比草地温度高，前者实测温度可超过 45 ℃,这种经常性、周期性高温，促使沙地植物采用与草地植物不同的适应策略，即沙地植物通过释放更多的异戊二烯或单萜来减少其可能遭的热胁迫或热伤害，这种长期适应策略，使沙地植物具有更高的萜类化合物释放潜力。 本文还调查了温带典型草地生态系统和退化草地生态系统的异戊二烯和单萜释放速率，结果表明典型草地的标准异戊二烯和单萜释放速率分别为0.50 μgC g-1 h-1和0.69 μgC g-1 h-1;退化草地的标准异戊二烯和单萜释放潜力分别为0.32 μgC g-1 h-1和1.59 μgC g-1 h-1。总的来说，温带草地的异戊二烯和单萜释放速率都比较低，尤其是典型草地。整个生长季，典型草地释放的异戊二烯和单萜分别为31.6 mgC•m-2　和 70.4 mgC•m-2;退化草地的异戊二烯和单萜释放量分别为20.8 mgC•m-2 和 168.8 mgC•m-2。退化草地萜类化合物总释放速率远高于典型草地，尤其是单萜释放能力。过度放牧引起的草地退化，通过改变植被种类组成，对温带草地的异戊二烯和单萜释放速率产生显著影响;总体而言，温带草地退化将会使草地释放更多萜类化合物。 在温带草地生态系统中，Clost as VOC相对其NPP而言很小，在环境PAR和温度高时,它的贡献率相对较大;Clost as VOC约占典型草地生态系统NEP的5.32 %，退化草地生态系统NEP的0.23 %。植物源VOC释放所损失的碳素相对草地生态系统NPP而言几乎可以忽略不计;但是，相对其NEP，Clost as VOC还是具有一定的相关性。虽然，草地生态系统Clost as VOC对NPP或NEP的贡献率较小，但考虑到全球尺度植物源VOC的巨大释放速率，它在碳循环中的贡献率仍然不容忽视;在某些特殊的生态系统中，仍可能扮演重要角色。

浑善达克沙地稀树疏林草地生态系统研究：生物量、生产力与生态恢复途径

荒漠化是困扰我国干旱和半干旱地区的重要环境问题，我国的四大沙地也是荒漠化最为严重的地区，从生态学角度提高对沙地的认识，探明其生物量和生产力对于了解沙地生态系统的结构和功能是非常有意义的。本研究以浑善达克沙地为例，对沙地的植被特点、生物量和生产力以及退化沙地的恢复策略进行了探讨。 通过对浑善达克沙地进行的植被调查发现，浑善达克沙地的稀树疏林草地景观由草本层、灌丛和榆树疏林构成，彼此呈相间排列、交替出现的格局。在丘间低地以地带性的草本层植物占优势，灌丛主要出现在沙丘和丘间低地的过渡地段，地下水位较高的地方。榆树（Ulmus pumila）是沙地的优势树种，占沙地乔木的95％，榆树疏林主要在固定沙地稀疏分布，占沙地榆树总量的84％。胸径10-25 cm的榆树占60％，幼树少见，这和放牧干扰有关。沙地的稀树疏林草地是不同于周围草原地带的隐域植被，带有明显的超地带性，同时因受地带性气候的影响，兼有地带性特征。 在碳循环方面，为了估测沙地植被的碳存储潜力，我们对浑善达克沙地的生物量进行了估测。整个沙地被分为六种生境，分别求算生物量，然后汇总。对于其中的榆树生物量建立了异速生长关系进行估算。结果表明，浑善达克沙地植被的平均生物量为21.30 Mg ha-1，其中丘间低地的贡献为65％。从生活型上看，草本层的贡献占优势，而榆树的贡献很小，仅为10％。平均地下/地上生物量的分配比例为2.9。与草地生态系统相比，浑善达克沙地的生物量要高出90％，表明沙地植被是草原地带内的隐域植被，特殊的土壤水热条件使它具有更高的碳存储潜力。沙地的大范围恢复将使该碳库通过碳获取得以实现。 净初级生产力（NPP）是生态系统功能的重要体现，它可以反映植被碳固定的能力。为了了解稀树疏林草地NPP的特点，我们对浑善达克沙地六种生境的NPP分别进行了求算，然后汇总。浑善达克沙地植被的平均NPP为11.06 Mg ha-1 yr-1，各生境的贡献不同，以丘间低地的贡献最大，达65％。生活型上看，草本层植物占优势，乔木的贡献仅为1.3％。地下/地上NPP为1.7，降水利用效率达10.9。浑善达克沙地稀树疏林草地的NPP比草地生态系统的平均值高59％。NPP的研究结果表明，沙地植被是和周围地带性草原植被存在显著差别的，这种高生产力将伴随沙地的恢复而实现，并有可能对沙地地区的碳循环产生影响。 为了探索荒漠化防治的新途径，在浑善达克沙地巴音胡舒嘎查开展了为期5年的退化草地恢复试验。结果表明，通过“以地养地”的模式可以使退化草地充分利用潜在恢复力进行自然恢复，并取得了明显效果。开辟高效地，种植高产玉米，在草地得到恢复的同时，牲畜仍然有充足的草料供应，牧民生活水平也有所提高。这种适合半干旱地区环境条件的恢复模式比在同样地区造林要有效和经济。据此，文章最后探讨了当前荒漠化防治中存在的误区和今后荒漠化防治应采取的策略。

克氏针茅(Stipa krylovii Roshev.)草原植物凋落物分解特性及其对环境变化的响应

凋落物的分解对生物地球化学循环起着重要的作用。本研究以位于中国北方农牧交错区（内蒙古多伦县）的半干旱克氏针茅(Stipa krylovii Roshev.)草原生态系统为背景，针对凋落物分解研究中的几个关键过程和问题进行了探讨。研究内容包括该生态系统中几种常见植物凋落物的分解速率，在分解过程中的养分动态，凋落物的混合效应（即非加性效应）及其机制，全球变化引起的土壤有效N、P和降水增加对凋落物分解的影响及其对草原生态系统碳储量的影响等。 通过对克氏针茅、糙隐子草(Cleistogenes squarrosa Trin.)、双齿葱(Allium bidentatum Fisch.)、野韭(Allium ramosum L. Sp. Pl.)和冰草(Agropyron cristatum Gaertn.)的叶、茎、根凋落物的研究发现，叶和茎的分解速率与凋落物初始N和P含量呈显著正相关，根系的分解速率与初始C/N比呈显著负相关。根系凋落物一般比叶和茎凋落物分解速率快，而且具有更快的养分周转速率。有些凋落物混合在一起分解后对分解速率产生了混合效应（混合凋落物的实际分解速率偏离于基于组分凋落物计算的预期分解速率），同时它们对养分（主要是N、P、Ca、Mg）的固定或释放被延后。混合效应发生与否以及混合效应的方向主要取决于组分凋落物的特点，而与凋落物多样性的高低没有明显的关系。混合凋落物对养分动态的负作用，能降低退化生态系统遭到干扰后养分的损失，而且养分释放时间的延后有助于某些植物更好地生长，有利于退化草原生态系统恢复过程中植被结构的改善。 由于凋落物的质量对分解速率的影响重大，凋落物的化学组成很可能对混合凋落物分解过程中的非加性效应起重要作用，然而迄今为止没有明确的结论。本文研究发现，具有不同初始N、P含量的凋落物（同一物种）混合在一起分解后产生了正的非加性效应，非加性效应的强弱与组分凋落物初始N、P含量的差异大小有关。而且，非加性效应与P含量差异大小的相关性比与N的强，这可能与研究地点土壤中P的含量相对更为缺乏有关。这一研究结果表明，凋落物的化学组成在有些情况下确实与非加性效应有关。 通过施肥和浇水试验，研究了全球变化引起的土壤有效N、P和水分的增加对具有不同生活型和化学组成的两种优势草原植物，即双齿葱和克氏针茅凋落物分解的直接影响。结果发现，向土壤中添加N肥或N、P复合肥，在短期内（100天）加速了这两种植物的分解速率。高质量凋落物（双齿葱）的分解更容易受到土壤水分条件的限制，而低质量凋落物（克氏针茅）的分解对土壤养分有效性更敏感。土壤中有效养分的增加会提高分解凋落物对养分的固定，有利于退化生态系统中养分的保持。双齿葱对内蒙古半干旱典型草原的碳循环和养分动态的贡献，比克氏针茅要更大。研究结果提示人们，对草原生态系统的碳循环和养分动态进行模拟时，需要对不同化学组成的凋落物分别加以考虑。 本文还探讨了土壤有效N长期增加后对两种优势草原植物（克氏针茅和冷蒿(Artemisia frigida Willd)）凋落物分解的综合影响，并进一步区分了由土壤有效N增加引起的凋落物质量改变的间接作用和土壤状况改变的直接作用。结果发现，土壤有效N长期增加后对凋落物的分解速率没有明显影响，但是显著加速了分解凋落物对养分的固定，这可能是由于凋落物质量对分解的促进作用和土壤状况对分解的抑制作用相互抵消的原因所造成的。向土壤中添加N素后，地上部植被生产力不可避免的会提高，尽管短期内会加速凋落物的分解速率，但从长远来看，对凋落物的分解速率不会产生影响，所以估计土壤有效N增加后最终会提高草原生态系统的净碳储量。 通过本研究，对中国北方农牧交错区（内蒙古多伦县）半干旱克氏针茅草原生态系统的养分循环过程有了初步的了解，验证了混合凋落物分解过程中非加性效应发生的可能性，并且证明组分凋落物的初始化学组成对这种非加性效应确实起着重要作用。本研究说明全球变化对不同植物种凋落物分解的影响是不同的，而且因研究时间长短的不同对凋落物分解的影响会发生变化。研究结果为草原生态系统碳和养分循环过程的模拟提供了必要的基础资料，为混合凋落物分解过程中非加性效应发生的可能机制提供了强有力的证据，对在全球变化背景下人们对草原生态系统碳和养分循环过程的认识和模拟有着重要的参考价值。

典型草原物候特征及其对生态系统功能的影响研究

植物物候反映的是植物(包括农作物)和环境(气候、水文、土壤条件)的周期性变化之间的相互关系。在气候变化背景下，植物物候已经发生了显著变化，并且对生态系统产生了重要影响。然而，目前的物候研究大多是针对木本植物，对于草本植物的研究则相对缺乏，而且草本植物的物候节律表现出较木本植物更为复杂的特征，不但受温度影响，亦受到水分因素的影响。 本研究利用内蒙古典型草原区克氏针茅草原建群种羊草和克氏针茅自1985～2003年19年的物候资料和气象数据，分析了物候特征及气候因子的变化趋势，探究了两种植物返青期和枯黄期的主导因子。结果表明，克氏针茅草原近20年来的气候发生了显著的变化，总体表现为温度升高、降水量降少、土壤水分含量减少。与以往物候研究结果不同的是，羊草和克氏针茅返青期在气候变暖的背景下却显著滞后。相关分析显示返青前期土壤水分是导致返青滞后的主要原因。对于枯黄期的相关分析同样显示水分因子是制约两种植物生长季结束的关键因子。在检验现有返青期和枯黄期物候模型对于典型草原适用性的基础上，本研究选择应用广泛、计算简便的CTM(Cumulative Temperature Model)模型作为改进返青期物候模型的基础，在其中加入了水分的影响，使得改进返青期物候模型可以很好的模拟典型草原植物返青期，模拟误差小于7天。同时，构建了考虑水分和温度共同影响的枯黄期模型。改进后的物候模型提高了DCTEM(Dynamic Chinese Terrestrial Ecosystems Model)模型的模拟精度。 基于耦合改进物候模块的DCTEM模型对影响生态系统NPP(Net Primary Productivity)、NEP(Net Ecosystem Productivity)和AET(Annual Evapotranspiration)的因子加以分析。结果显示，降水量是影响克氏针茅草原生态系统功能的主要因子，其对于NPP、NEP、AET以及土壤异养呼吸等均有不同程度的影响。其次，生长季长度变化对于克氏针茅生态系统功能呈现出显著的影响作用，其影响程度仅次于降水量。 为了量化在实际的气象条件下单位生长季长度变化所引起生态系统NPP、NEP和AET的变化幅度，设置了三个引起生长季长度变化的物候模拟情景(动态枯黄情景、动态返青情景、动态起止情景)以及对照情景。研究结果显示，不同情景下植物生长季长度变化对于生态系统功能有着不同程度的影响。动态枯黄情景下由于草原枯黄期使整个生长季每延长一天NEP增加3.11％，NPP为0.34％，对于AET的影响最小为0.06％;动态返青情景下，由于草原返青期波动使得整个生长季延长一天则NEP增加1.54%，NPP为0.15％，对于AET的影响最小为0.01％;在动态起止情景下，生长季延长一天则NEP增加3.37%，NPP为0.39％，对于AET的影响最小为0.06％。总体而言，由于枯黄期引起的生长季变化对生态系统功能影响程度比由于返青期引起的程度高。此外，不同的生态系统功能要素对于物候变化的影响程度也有所不同。在几种模拟情景下，NEP受到生长季变化的影响最大，其次为NPP，AET受物候变化影响最小。

改变土壤碳输入对我国北方温带草原土壤碳、氮库和碳、氮循环的影响

土壤是陆地生态系统最重要的碳库和氮库，在陆地生态系统碳氮循环过程中起着举足轻重的作用。植物光合作用产物向地下的分配(根系分泌、细胞凋亡和根系分解)和地上凋落物分解是土壤碳两个最主要的来源，关于这两种碳输入途径和量如何影响陆地生态系统土壤碳氮库和循环的有限理解，限制了我们预测陆地生态系统碳循环对于全球气候变化的响应和反馈。本实验以我国北方广泛分布的温带典型草原生态系统为研究对象，通过添加和去除地面凋落物以及割草改变植物对土壤的碳输入途径和量，观察温带草原土壤有机碳库和氮库各个组分(微生物生物量碳氮、活性碳、可溶性碳、轻重组碳、总有机碳、总有机氮)及碳氮循环(微生物呼吸、土壤呼吸、生态系统气体交换、净氮矿化)的变化，来区分和量化凋落物分解和根系分泌、分解这两种碳输入方式对草原土壤碳库和循环的相对贡献和影响，有助于揭示在全球变化背景下温带典型草原生态系统碳库和碳循环的响应机制，并为预测其未来变化动态以及对全球气候变化的反馈提供参数估计和模型校正和验证。 我们对各处理样方土壤温度、土壤水分以及土壤呼吸进行了一个生长季的测定，结果表明，添加和去除凋落物都使得土壤呼吸与对照相比都有所增加，但是都没有达到显著水平;而无论地上凋落物的量添加、去除还是不变，植物光合作用对土壤呼吸的影响则比较显著，去除植物显著地降低了土壤呼吸。这说明处理一年后，植物对土壤呼吸起主导作用，而凋落物处理对土壤呼吸的影响还需要更长时间的观察。此外，生物量的结果也表明了地上凋落物的改变会影响到植物生长，从而影响到植物对土壤的碳输入，但是都没有达到显著水平，仍然需要我们进行更长时间的处理和观察。 通过在生长季前期、中期、后期进行了三次氮矿化的测定，以及在生长季后期进行了一次微生物生物量测定，我们得出以下结论：添加凋落物加快了净氮矿化速率，但是没有达到显著影响水平;同样，去除凋落物降低了净氮矿化速率，也没有达到显著水平。此外，添加凋落物处理微生物生物量碳、氮都有所增加，与对照相比没有显著差异，但是显著高于另外几个处理。 通过本研究，我们对于根系分泌和地上凋落物分解这两种碳氮输入途径对我国温带草原碳氮循环的影响有了初步的了解。与森林生态系统相比，由于草原生态系统土壤碳氮含量低，土壤碳库和转化对碳输入途径和量的变化更为敏感，在处理第一年，土壤微生物量和土壤呼吸都发生了显著的变化。然而，由于改变土壤碳库是长时间日积月累的结果，草原生态系统某些指标对碳输入途径和量的变化的响应还需要较长时间才能显示出来，例如添加和去除凋落物虽然使得土壤呼吸值有所增加，但是还都没有达到显著的水平;另外碳输入途径和量的变化对草原生态系统氮矿化还没有产生显著的影响。这些都需要更长时间的处理和观测来验证。