Spatial and temporal relationships between precipitation and ANPP of four types of grasslands in northern China

Precipitation is considered to be the primary resource limiting terrestrial biological activity in water-limited regions. Its overriding effect on the production of grassland is complex. In this paper, field data of 48 sites (including temperate meadow steppe,temperate steppe, temperate desert steppe and alpine meadow) were gathered from 31 published papers and monographs to analyze the relationship between above-ground net primary productivity (ANPP) and precipitation by the method of regression analysis. The results indicated that there was a great difference between spatial pattern and temporal pattern by which precipitation influenced grassland ANPP. Mean annual precipitation (MAP) was the main factor determining spatial distribution of grassland ANPP (r~2 = 0.61,P ＜ 0.01); while temporally, no significant relationship was found between the variance of AN PP and inter-annual precipitation for the four types of grassland. However, after dividing annual precipitation into monthly value and taking time lag effect into account, the study found significant relationships between ANPP and precipitation. For the temperate meadow steppe, the key variable determining inter-annual change of ANPP was last August-May precipitation (r~2= 0.47, P = 0.01); for the temperate steppe, the key variable was July precipitation (r~2 = 0.36, P = 0.02); for the temperate desert steppe, the key variable was April-June precipitation (r~2 = 0.51, P ＜0.01); for the alpine meadow, the key variable was last September-May precipitation (r~2 = 0.29, P ＜ 0.05). In comparison with analogous research, the study demonstrated that the key factor determining inter-annual changes of grassland ANPP was the cumulative precipitation in certain periods of that year or the previous year.

Spatial and temporal patterns of aboveground net primary productivity (ANPP) along two freshwater-estuarine transects in the Florida Coastal Everglades

We present here a 4-year dataset (2001–2004) on the spatial and temporal patterns of aboveground net primary production (ANPP) by dominant primary producers (sawgrass, periphyton, mangroves, and seagrasses) along two transects in the oligotrophic Florida Everglades coastal landscape. The 17 sites of the Florida Coastal Everglades Long Term Ecological Research (FCE LTER) program are located along fresh-estuarine gradients in Shark River Slough (SRS) and Taylor River/C-111/Florida Bay (TS/Ph) basins that drain the western and southern Everglades, respectively. Within the SRS basin, sawgrass and periphyton ANPP did not differ significantly among sites but mangrove ANPP was highest at the site nearest the Gulf of Mexico. In the southern Everglades transect, there was a productivity peak in sawgrass and periphyton at the upper estuarine ecotone within Taylor River but no trends were observed in the C-111 Basin for either primary producer. Over the 4 years, average sawgrass ANPP in both basins ranged from 255 to 606 g m−2 year−1. Average periphyton productivity at SRS and TS/Ph was 17–68 g C m−2 year−1 and 342–10371 g C m−2 year−1, respectively. Mangrove productivity ranged from 340 g m−2 year−1 at Taylor River to 2208 g m−2 year−1 at the lower estuarine Shark River site. Average Thalassia testudinum productivity ranged from 91 to 396 g m−2 year−1 and was 4-fold greater at the site nearest the Gulf of Mexico than in eastern Florida Bay. There were no differences in periphyton productivity at Florida Bay. Interannual comparisons revealed no significant differences within each primary producer at either SRS or TS/Ph with the exception of sawgrass at SRS and the C−111 Basin. Future research will address difficulties in assessing and comparing ANPP of different primary producers along gradients as well as the significance of belowground production to the total productivity of this ecosystem.

Grassland productivity limited by multiple nutrients

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)2,3, the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized4,​5,​6,​7,​8. However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Anthropogenic nitrogen deposition predicts local grassland primary production worldwide

Humans dominate many important Earth system processes including the nitrogen (N) cycle. Atmospheric N deposition affects fundamental processes such as carbon cycling, climate regulation, and biodiversity, and could result in changes to fundamental Earth system processes such as primary production. Both modelling and experimentation have suggested a role for anthropogenically altered N deposition in increasing productivity, nevertheless, current understanding of the relative strength of N deposition with respect to other controls on production such as edaphic conditions and climate is limited. Here we use an international multiscale data set to show that atmospheric N deposition is positively correlated to aboveground net primary production (ANPP) observed at the 1-m2 level across a wide range of herbaceous ecosystems. N deposition was a better predictor than climatic drivers and local soil conditions, explaining 16% of observed variation in ANPP globally with an increase of 1 kg N·ha-1·yr-1 increasing ANPP by 3%. Soil pH explained 8% of observed variation in ANPP while climatic drivers showed no significant relationship. Our results illustrate that the incorporation of global N deposition patterns in Earth system models are likely to substantially improve estimates of primary production in herbaceous systems. In herbaceous systems across the world, humans appear to be partially driving local ANPP through impacts on the N cycle.

内蒙古锡林河流域草原植物和群落热值的时空变异研究

第一部分：内蒙古锡林河流域草原植物种群和群落热值的时空变异研究 热值的研究是评价生态系统能量固定、传输和转化的基础，也是评价植物光合作用效率和植物营养值的有用参数。同种植物热值会随着植物部位、光照、养分条件、季节、土壤类型和气候条件的不同而发生变化。不同的种类和类群之间热值也存在差异。本项研究以内蒙古锡林河流域中段草原植物群落为对象，研究了植物种群和群落热值的时空变异规律。 对内蒙古羊草草原群落不同植物种群热值的时问动态研究结果表明，42种植物地上部分的热值在13.16土1.14 kJ.g-l和18.14土0.53 kJ.g-1之间变动，所有物种的平均热值为16.90土0,84 kJ.g-1，种间变异系数4.9%。小叶锦鸡儿(Caraganamicrophylla)具有最高的热值。禾草的平均热值高于杂草。根据生活型和生长型，草本物种被进一步分组，热值从高到低的排列顺序为：高禾草>豆科植物>矮禾草>其余杂草>半灌木>一二年生植物。 主要植物种群地下部分热值的分布范围为15.05-16.41 kJ.g-1。其中根茎型草地下部分热值较高。不同种类植物地下部分热值差异并不与地上部分一致。根茎型禾草地上、地下部分热值差异较小，而须根型植物差异较大。不同种群的植物地上部分热值随植物物候期的不同而波动，其变化规律是与植物种群本身的生物学特性相联系的。不同植物种群热值的年际波动规律有所不同，羊草(Leymuschinensis)、大针茅(Stipa grandis)和洽草(Koeloria cristata)的年际热值波动相关显著，但与生长季降水量和生长季累积日照时数之间无明显相关性。在某种程度上，植物热值的种内变化反映了植物生长状况的差异。 42种植物的热值和它们在群落中的相对生物量存在显著正相关关系。表现为优势种(17.74 kJ.g-1)>伴生种(17.24 kJ．g-l)>偶见种(16.65 kJ.g-1)。高热值的植物更具竞争力，在群落中通常占据优势地位，而低热值的植物竞争力通常较弱，构成草原群落的伴生种或偶见种。 以内蒙古锡林河流域3个草原群落类型（羊草典型草原，大针茅典型草原，羊草草甸草原）的放牧退化梯度系列（包括未退化，轻度退化，中度退化和重度退化4个强度）为研究对象，对主要植物种群和群落热值随草原类型和退化梯度的空间变异规律及热值与其他群落和土壤性质的相关性进行了研究。 结果表明，研究区出现的60个植物种平均热值为17.25土0.92 kJ.g-1，变异系数5.4%．热值大于18.00 kJ.g-1的高能植物包括3种优势高禾草(羊草、大针茅和羽茅(A. sibiricum))和一些有毒植物，热值小于17.00 kJ.g-1的低能值植物包括多数一年生杂草;热值在17.00-18.00 kJ.g-1之间的中能值植物包括大多数多年生杂草和矮禾草。 按照生活型分类，灌木的热值最高，多年生禾草显著高于一二年生植物，半灌木和多年生杂草介于二者之间。按照水分生态类型分类，旱生植物、中旱生、旱中生和中生植物之间在热值上没有明显差异。不同科之间热值存在显著差异，禾本科、豆科、菊科植物热值较高，藜科植物平均热值最低。 二因素方差分析结果表明，主要优势物种热值在不同草原类型之间存在显著差异，表现为羊草草甸草原>羊草典型草原>大针茅典型草原。对于大多数优势禾草，热值没有随退化梯度发生明显变化，洽草(K. cristata)、冰草(A．ctistatum)和所有优势杂草随退化程度的增强热值趋于下降。对于大多数优势物种，热值随不同草原类型的空间变异大于放牧退化所导致的空间变异。 不同草原类型的群落热值为羊草草甸草原>羊草典型草原>大针茅典型草原，群落平均热值表现出随退化强度的增加而下降的趋势，这主要归因于沿退化梯度不同物种构成比例的变化，即随退化程度的加剧，高能值植物在群落中的比例下降。其次是特定物种热值随退化梯度的变化。在同一草原区，放牧对群落热值的影响大于立地条件之间的差异。 群落和主要物种热值均表现出与某些群落特征和土壤性质的相关性。 关键词：内蒙古，锡林河流域，羊草草原;物种和群落热值，时空变异，退化梯度，草原类型，土壤性质 第二部分内蒙古羊草草原17年刈割演替过程中功能群组成动态及其对群落净初级生产力稳定性的影响 基于17年的野外实验数据，研究了内蒙古羊草草原群落刈割演替过程中的功能群组成动态，探索功能群组成变化与群落净初级生产力(ANPP)之间的关系，分析结构参数怎样影响功能参数。结果显示：在17年的割草演替过程中，群落的结构与功能均发生了变化。随着羊草群落刈割演替的进行，群落的功能群组成发生了显著变化，根茎禾草在群落中的优势地位相继被一二年生植物，高丛生禾草，矮丛生禾草所取代。到17年末，群落变成根茎禾草，矮丛生禾草，高丛生禾草共同建群的群落。在对照群落中ANPP与年降水量显著相关，但在刈割群落中二者则不相关。年降水量解释对照群落ANPP变异的62%，而连年的刈割干扰则是刈割群落中ANPP动态的主要驱动因子。群落净初级生产则显出对刈割干扰的抵抗能力，在刈割干扰的前几年，依靠群落内功能群组成的不断调节，保持相对稳定的水平，当刈割进行5年之后，群落结构的变化积累到一定程度，净初级生产迅速下降到一个较低的水平，此后依靠群落结构的不断调节来维持这一功能水平。因此，群落结构是以渐变的方式改变的，而群落功能的下降则是以跃变的形式完成的。群落依赖于结构的不断调整来保持功能的相对稳定，但结构变化到一定程度也会导致功能的衰退。 关键词：内蒙古，羊草草原;刈割演替;功能群组成;净初级生产;群落;稳定性

内蒙古典型草原地上净初级生产力估测及模型模拟的研究

地上净初级生产力（ANPP）是陆地生态系统碳循环的重要组成部分，但由于估测ANPP的方法不同使得对ANPP的估测值存在很大的不确定性。本文采用3种方法（群落中所有种群当年最大地上生物量之和（ANPPC1）、当年群落最大地上生物量（ANPPC2）以及每年固定日期（8月30日）的地上生物量（ANPPC3））在种群、功能群和群落水平上分别对连续19年（1980～1998）的内蒙古羊草和大针茅草原生态系统功能（如ANPP、植物多样性和水分利用效率（WUE））的动态变化进行了比较分析，同时探讨了不同估测方法下气候和放牧对ANPP的影响，在此基础上利用DNDC模型进行了ANPP的模拟和敏感性分析研究，主要结果包括： 在羊草和大针茅草原群落中，不同主要植物种群或功能群多年平均地上最大生物量出现的时间不同，而同一植物种群或功能群的年地上最大生物量出现的时间存在年际间的变化。采用当年最大地上生物量和8月30日固定日期的地上生物量作为群落ANPP的这两种方法高估了建群种或禾草功能群在群落中的作用。 羊草草原群落多年平均ANPPC1、ANPPC2和 ANPPC3分别是257.5、190.1和166.0 g.m-2;相应的大针茅草原多年平均ANPPC1、ANPPC2和 ANPPC3分别是180.4、132.8和122.5 g.m-2。就群落生产力而言，后两种常用的方法二和方法三分别低估了草原群落ANPP 14.2%～40.0%和15.5～59.0%。本文研究表明尽管ANPPC1与ANPPC2和ANPPC3之间存在显著的差异，但二者之间存在极显著的相关性:羊草草原的ANPPC1= 59.587+1.061×ANPPC2（r2＝0.865, p＜0.001），ANPPC1= 92.329+1.017×ANPPC3（r2＝0.569, p＜0.001）;大针茅草原的ANPPC1= 32.918+1.114×ANPPC2（r2＝0.814, p＜0.001），ANPPC1= 76.120+0.875×ANPPC3（r2＝0.499, p＝0.001）。 种群、功能群和群落地上净初级生产力与气候因子间的关系因不同的估测方法而异。羊草草原的建群种羊草种群仅ANPPS3与8月和11月份的平均温度间存在显著相关性，而大针茅草原群落的建群种大针茅种群的ANPPS1和ANPPS2与3月份平均最高气温间呈负相关关系。在羊草草原群落，杂类草功能群ANPPF1和ANPPF2与3月份气温呈负相关关系;而灌木半灌木功能群ANPPF1和ANPPF2与3月份降水呈负相关关系。在大针茅草原群落，禾草功能群ANPPF1与5月份最高平均气温呈显著负相关关系。羊草草原群落ANPPC1和ANPPC3分别与11月份最低温度和平均温度存在显著负相关关系;大针茅草原群落ANPPC1与2和6月份月降水量间相关性显著，ANPP2与5月和11月份月平均最低温度呈显著负相关关系，而4～9月份，1和4月份平均最低温度对群落ANPPC3起决定作用。 在羊草和大针茅草原群落，由方法一得到的群落水分利用效率、Shannon植物多样性指数和均匀度指数与方法二或方法三得到的相应的指数间存在显著的差异，方法二和方法三得到的值间差异不显著。群落地上净初级生产力与相应估测方法的植物Shannon多样性指数和均匀度指数之间相关性不显著。 放牧条件下羊草或大针茅草原群落中的建群种羊草和大针茅在群落中的相对地上生物量较围栏内的相应植物种群的降低，而糙隐子草种群在群落中的比例上升。不同的放牧管理条件下，群落中植物种群地上现存量的季节动态发生变化。群落的植物组成及植物种群地上生物量在群落中总生物量的比例发生了明显的变化。利用遥感来估测地上净初级生产力时，分别低估了羊草和大针茅草原ANPP 52％和27％。 DNDC模型可以很好地模拟内蒙古典型草原生态系统的地上生物量，通过敏感性分析表明降水是草原植物生长的主要限制因子，在降水量增加或降低至日降水的30%时，模拟的地上生物量显著地高于或低于实测地上生物量值。在多年平均降水量为347mm的情况下，随着土壤粘粒含量的增加，地上生物量逐渐降低，与北美草原一致。

施肥和降水增加对半干旱草地生产力和温室气体排放的影响

人类活动能显著地改变生态系统的属性和功能。这些变化不仅发生在容易观察的局部层面上，如土壤沙化、植被退化等;还能更深刻地影响到微生物控制的温室气体释放，进而影响大尺度上气候的变化。本论文根据2 年田间试验结果报告了人类干扰（草地退耕和施肥）和降水增加30%对中国北部农牧交错区（内蒙古多伦县）半干旱草地生态系统生产力和温室气体排放的影响。 本研究发现氮是影响该区草地和弃耕地生产力的最重要因子，即使低量的氮肥应用(5 g N y-1)也能使该区草地地上净初级生产力（ANPP）增加36-46%。然而氮对草地生产力影响主要集中在地上部分，对地下净初级生产力（BNPP） 影响比较微弱。单独使用磷肥对ANPP 和BNPP 均无显著影响，氮磷混施对ANPP 有显著正交互作用。增加的降水对草地和弃耕地ANPP 和BNPP 有弱的正效应，增加30%降水使草地对照ANPP 增加9-25%，BNPP 增加12%-17%，弃耕地对照ANPP 增加7%-37%，BNPP 增加36%-37% 。草地和弃耕地对照ANPP 并无显著差别，由于高于弃耕地1-3 倍的BNPP，总体上草地比弃耕地有更高的固碳能力。这个结果也说明草地能更有效地把植物地上部分固定的碳转移到地下，这与草地长期发育积累的丰富地下根系和高根冠比物种组成有关。 水肥处理及土地利用方式也深刻改变了温室气体排放。温带草地和弃耕地CO2 排放（系统呼吸：Re）存在显著季节变化，温度、水分和地上生物量是控制这些变化的主要因子。在气候温暖的6-9 月份，生物量和土壤水分是决定系统呼吸季节变化主要因子，而与温度无明显相关。随着氮肥使用，草地和弃耕地呼吸速率加大，其温度敏感系数Q10 也明显升高，这主要与氮肥使用后增加的地上生物量有关。由于水分促进作用和生物量的弱增长，增加的降水也加大了草地和弃耕地CO2 排放。低的地下生物量和有机碳氮并没有导致弃耕地低的CO2 排放，显示了农垦后土壤结构破坏导致的有机碳分解增加不会在农业弃耕后短期内（5-6 年）停止。 除了对生物量的不敏感性外，中国北部半干旱生态系统N2O 排放表现出与CO2 排放类似的变化规律：水分和温度是N2O 季节变化的主导因子;氮肥使用和降水增加都增加了N2O 排放;N2O 排放在草地对照和弃耕地对照之间没有显著差别。更少的根系对无机氮的竞争导致了弃耕地比草地有更高的N2O 释放因子（单位氮添加所引起N2O 释放）。低的土壤水分（<70% WFPS ）和无机氮都是限制反硝化发生的重要条件，因此硝化可能是草地和弃耕地N2O 的最主要来源。半干旱草地生态系统N2O 排放与CO2 排放之间存在着显著线性关系。 中国北部半干旱草地和弃耕地都起着CH4 汇的生态功能。水分是控制CH4 吸收季节变异的主导变量，土壤空隙水含量（WFPS）与CH4 指数递减方程能解释其变异的64%-83% 。增加的降水导致了土壤水分增加，从而引起甲烷氧化菌所需基质（CH4 和O2）扩散受阻，减小了土壤CH4 吸收功能。施肥并没有抑制CH4 吸收，相反由于施肥后植物速生对土壤水的抽干作用，施肥增加了植物速生期时的CH4 吸收。CH4 吸收对农垦引起的土壤微生物生境结构破坏非常敏感，在农业措施停止5-6 年后的弃耕地，CH4 吸收仍比草地低24%。 本研究初步分析了影响中国北部农牧交错区草地生态系统生产力及温室气体排放的自然环境因子和人为因子，给出了它们之间的定性、定量关系，为科学地管理中国北部草地，充分发挥其生态和经济效益功能提供了有效信息。

内蒙古多伦典型草原几种植物叶片性状与生物量及降雨量关系的初步研究

植物性状是植物与环境长期互作过程中表现出的内在或外部的特征。目前植物生态学界关于植物性状的研究，主要包括以下三个方面的研究内容：植物性状与环境因子间的关系，反映植物性状对环境变化的响应或适应机制；植物性状与生态系统功能的关系，以更好地揭示生态系统功能的内在驱动机制；植物性状间的相关关系和消长对策，反映植物性状的协同进化机制。 本试验所在地区内蒙古多伦十三里滩为典型克氏针茅草原，水分为当地生态系统的主要限制因素。我们在当地的围封样地内，采用人工模拟生长季降雨变化梯度的方法，研究了内蒙古典型草原几种常见植物叶片功能性状对模拟降雨量梯度的响应、性状之间的协变关系以及植物性状与地上部分生物量间的关系。研究工作在2005-2006年期间进行。观测的叶片性状包括叶片干物质含量(LDMC)，比叶面积(SLA)和叶片氮含量(LNCmass, LNCarea)，以地上部分生物量作为年净初级生产力(ANPP)估计，降雨量梯度模拟了两个水平，即对照(接收正常降雨)和增雨。主要结论如下：1) 叶片性状在降雨量梯度上的变化趋势在物种水平上表现出多样化的特点，性状的变化趋势是否有显著的格局取决于物种或特定性状。2) 叶片性状对水分梯度的响应在功能群水平上表现出各自的特点，Grass功能群对水分梯度的响应敏感性较低，Forb居中，Legume表现出较强的敏感性，表明Grass功能群对水分的耐受性比较强，这在一定程度上解释了当地克氏针茅草原上目前以禾草类为主的现状，对物种之间的替代有一定的指示性。而群落水平上探讨性状在环境梯度上的变异趋势，研究结论存在较大的不确定性，主要是难以区分环境和系统发育背景对植物性状的影响。3)叶片功能性状间的协变关系，不同物种因其自身发生背景不同，在不同环境背景下总体表现出一致的变化规律，但也存在例外。相对而言，对功能性状间协变关系的研究在物种水平或功能群水平上更有意义，而在群落水平上所得结论，可能会掩盖了群落内部不同物种或不同功能群间的信息。4)叶片功能性状与草原地上部生物量的关系在功能群水平上呈显著正相关关系，再次验证了土壤含水量的变化同土壤氮元素的动态，共同影响了植物的氮利用策略，进一步影响了初级生产力等和氮元素直接相关植物性状参数。在群落水平上叶片功能性状与生物量的之间并没有表现出可遵循的规律性，说明叶片功能性状对生物量的作用更多地局限在群落内部的组织层次上发挥作用。5)本研究的虽然从不同的组织层次上探讨了叶片功能性状对环境、性状之间的协变关系以及与地上部分生物量间的关系，由于植物性状变化幅度可能与环境梯度密切相关，如果模拟环境梯度不够大，可能会影响结论的普遍性。此外，本次野外控制试验时间较短，数据量偏少，本文的结论尚需要进一步验证。

内蒙古温带草原群落结构和生态系统功能对火烧、氮素添加和地形的响应

我国北方温带草原是欧亚大陆草原生物群区的重要组成部分，对于区域和全球的碳循环和平衡起着重要的作用。频繁的自然或人为干扰能够改变草原生态系统的群落结构和生态系统功能，从而影响生态系统为人类提供的产品和服务。本研究选取位于内蒙古多伦县的半干旱温带草原，研究火烧、氮素添加和地形以及它们的综合作用对该地区植物生产力、植物多样性、盖度和土壤呼吸的影响；另外，我们比较研究了由于地形因素而存在于草原地区的林地群落与其邻近草地的碳氮库和循环；旨在探讨我国北方温带草原地区人为干扰对草原生态系统结构与功能的影响以及该地区林地和草地碳氮库和循环的差异机理，以期为模型模拟本地区的生态系统碳循环提供理论依据和数据支持，具体研究结果如下： 1. 2006–2008 年，通过研究植物多样性和盖度对地形、火烧和氮素添加及其交互作用的响应，结果表明：半干旱草原植物物种数、香农威纳指数、均一性指数和盖度均表现出显著的年际变化。坡下的物种数、香农威纳指数和均一性指数均低于坡上。坡下较高的羊草(Leymus chinensis)、冰草(Artemisia frigida)、唐松草(Thalictrum petaloideum)和冷蒿(Agropyron cristatum)的盖度导致坡下的群落总盖度、禾本科草和非禾本科草盖度分别比坡上高22.5%、9.6%和13.2%。春季火烧提高了物种数、香农威纳指数和均一性指数。火烧对群落总盖度影响较小是由于火烧后非禾本科草冷蒿盖度的降低抵消了禾本科草羊草、冰草和针茅(Stipa kryroii)盖度的增加。施氮肥后物种数、香农威纳指数和均一性指数均降低。禾本科草羊草、冰草和针茅以及非禾本科草唐松草盖度的增加导致施肥后群落总盖度、禾本科草和非禾本科草的盖度分别增加了23.6%、35.1%和21.2%。火烧对禾本科草和非禾本科草盖度的作用受地形和氮素添加的影响。地形、火烧和氮素添加对植物盖度的影响主要受土壤水分调控。 2. 2005–2008 年，通过研究净初级生产力(NPP)对火烧、氮素添加和地形及其交互作用的响应，结果表明：半干旱草原的NPP 具有显著的年际变化。火烧后地上净初级生产力(ANPP)、地下净初级生产力(BNPP)和BNPP/ANPP 分别增加了12.8%、22.2%和14.9%。ANPP 的提高是由于火烧后禾本科植物(主要是羊草、冰草和针茅)生物量的增加。与之相反，火烧降低了非禾本科草，特别是冷蒿的生物量。氮素添加提高了ANPP (54.8%) ，对BNPP 没有影响，导致施氮肥后BNPP/ANPP 显著降低(33.4%)。禾本科草羊草、冰草和针茅以及非禾本科草唐松草生物量的增加，是氮素添加提高ANPP 的主要原因。坡下的ANPP 和BNPP 分别比坡上高14.1%和8.2%，但地形对BNPP/ANPP 没有影响。坡下ANPP 的提高主要是由于坡下禾本科草羊草、冰草以及非禾本科草唐松草、冷蒿的生物量高于坡上。氮素添加和地形影响ANPP 和BNPP/ANPP 对火烧的响应。火烧、氮素添加和地形对NPP 和植物碳分配39%–75%的综合效应可由这三个因素的简单加和效应来解释。 3. 通过研究2005 和2006 年生长季内土壤呼吸对地形、火烧和氮素添加的响应，结果表明：坡下的季节平均土壤呼吸比坡上高6.0%。春季火烧在整个生长季内促进土壤呼吸，平均增幅达23.8%。另外，火烧对土壤呼吸的效应受到季节和地形的影响。施用氮肥增加了11.4% 的土壤呼吸。火烧和地形对土壤呼吸的影响主要受土壤水分和植物生长的调控；而施氮肥后土壤呼吸的增加，主要是由于氮素添加促进植物生长后根系活性和呼吸的提高。 4. 2006–2007 年，通过对林地群落及其邻近草原生态系统土壤温度、土壤水分、土壤机械组成、地上和地下生物量、凋落物现存量、土壤碳氮储量、土壤呼吸、氮矿化和土壤微生物生物量的比较研究，结果表明：林地的土壤温度比草地低5°C，而其土壤水分却比草地高3.1%(绝对差异)。尽管林地(11,928.1 g m–2)和草地(11,362.2 g m–2)的土壤碳储量差异不显著，由于林地较高的植物生产力导致其碳储量高于草地。与草地相比，林地具有较高的凋落物现存量及碳氮含量、土壤无机氮含量、矿化氮的累积量、微生物生物量碳、微生物生物量氮、土壤呼吸和微生物呼吸。草地和林地的氮矿化速率没有显著差异。由地形因素引起的水分差异对于调控林地和草地生态系统碳氮库和循环(土壤碳氮储量、BNPP、矿化氮的累积量)具有重要作用。林地与草地生态系统碳储量的差异影响了我国北方草原地区碳的评估。

半干旱区沙质草地关键生态系统过程对水肥添加的响应

制约半干旱区生态系统恢复的主要因素是什么？一直是生态学界争论的问题，水分或养分或二者的联合限制是现存的3种观点，但是利用野外实验结合生态化学计量比，分析和判断半干旱区自然生态系统生产力恢复的限制因子还未见报道。 本论文以科尔沁沙地东南缘退耕后自然恢复的沙质草地为研究对象，通过析因设计的水、氮、磷添加实验，研究与碳循环、氮循环、磷循环等有关的生态系统关键过程，及其对人为干扰（灌溉和施肥）及全球变化（气候变化和氮沉降）的响应。从2004年到2005年采用3因素2水平析因设计进行水（0, 80mm）、氮（0, 20 g N m-2 yr-1）、磷（0, 10 g P2O5 m-2 yr-1）添加实验。2005年增加了0、2.5、5、7.5、10、30、40 g N m-2 yr-1的氮梯度添加实验。通过两年的实验，得出主要结论如下： 1）该沙质草地生态系统生产力主要受氮素养分的限制，水分与磷素并不是主要限制因素。施氮量达到7.5 g N m-2 yr-1时生物量明显提高，当氮素添加量达到40 g N m-2 yr-1时，生产力最高为1607.3 g m-2，但是并未找出氮肥添加量的上限。禾本科生物量与Shannon-Wiener多样性指数间呈指数负相关，拟合方程为：y = 1318.3e-0.2421x（R2 = 0.6887）。 2）添加水增加了土壤CO2排放速率，而施氮肥的影响并不明显。在较干旱的年份（降水量低于450 mm）干旱期（4月15日-6月15日）添加磷肥明显抑制了土壤呼吸，而在较湿润的年份（降水量高于450 mm）干旱期添加磷肥则显著增加了土壤呼吸。土壤呼吸与表层0-10 cm土壤含水量存在显著的正相关关系；土壤呼吸与土壤温度之间呈显著的指数正相关（R2 = 0.6798），拟合方程为：y = 0.1832e0.1299x。 3）氮素添加对半干旱区沙质草地生态系统氮有效性具有明显的增强作用，可提高土壤氮的矿化速率。该沙质草生态系统土壤有效氮主要由NO3--N组成，并具有明显的季节变化，NH4+-N和NO3--N含量在表层0-10 cm有高于10-20 cm的趋势。植物地上部分和凋落物的C:N比极低。土壤全量的C:N比明显受添加水的影响，与氮、磷添加无关。 4）添加磷素改善了半干旱区沙质草地生态系统土壤磷的有效性，添加氮和水则没有影响。0-10 cm层土壤相对有效磷有高于10-20 cm层的趋势，这说明土壤中的有效磷淋溶作用不强或者是植物主要利用了表土层10 cm以下的土壤有效磷。第一次提出P:N比的概念，利用相对有效磷和相对矿质氮的比值作为土壤相对P:N比，相对P:N比与生物量、ANPP、非禾本科ANPP之间呈显著的线性相关关系。相对P:N比与生物量的线性负相关程度最高，拟合方程为：y = -52.333x + 1356.2（R2 = 0.7263）。 5）水、肥添加对该沙质草地土壤含水量的影响并不显著，不同处理之间以及相同处理的不同层次之间土壤含水量的差异主要源自土壤的空间异质性。根据2005年4-7月的土壤含水量的方差分析结果，可以将表层0-10 cm划分为多变层，10-30 cm划分为过渡层，30-100 cm划分为稳变层。单独添加磷肥或水磷结合可以显著降低白草的蒸腾速率，而且添加磷肥可以抵消添加氮肥所导致的蒸腾速率增加。 6）添加水提高了物种丰富度和均匀度，而施氮肥降低了物种丰富度和均匀度，施磷肥也相应地增加了物种丰富度和均匀度，但幅度不大。本项研究的结果不支持单调上升格局，多样性与生产力呈负相关关系，单调下降和单峰关系两种格局都存在。10-20 cm土层相对矿质氮与物种丰富度、Shannon-Wiener指数、Shannon-Wiener均匀度指数均呈显著的负相关关系。物种丰富度与土壤相对矿质氮之间负相关关系最显著，可以用幂函数进行拟合方程：y = 348.58x-2.1236（n = 8; R2 = 0.8576）。 综合以上的研究结果，建议适量施用氮肥，加速退化沙质草地生态系统生产力的恢复；干旱年份辅以磷肥，以降低温室气体的排放速率。

降雨量变化对科尔沁沙地东南部樟子松人工林主要生态过程的影响

水分条件是影响植物生长最主要的限制因子，降雨量变化作为全球变化的一个重要组成部分，其对干旱半干旱区陆地生态系统的影响甚至超过CO2浓度和温度的升高以及它们的共同作用对生态系统的影响。樟子松人工林是科尔沁沙地东南部主要的防风固沙林类型，研究未来降雨量变化对会对樟子松人工林产生怎样的影响，对樟子松人工林的可持续经营和科学管理有重要意义。本研究以樟子松人工林为研究对象，通过搭建遮雨棚，铺设灌溉设施，野外原状样地模拟三个降雨量梯度：降雨量减少30%、天然降雨量和降雨量增加30%，从樟子松人工林下土壤生态系统、樟子松针叶生理特性、樟子松的生长和林下植被结构与生产力三个角度研究降雨量变化对樟子松人工林主要生态过程的影响，主要结论如下： （1）以土壤矿质N含量为土壤N有效性的指标，2007年的数据表明降雨量减少时土壤N有效性显著升高，降雨量增加时土壤N有效性显著降低，出现了“水、N有效性的不同时性”，即土壤水分有效性高时N有效性低，而N有效性高时水分有效性低，这可能是该地区植物生长的主要限制因子，而不是简单的水分限制或者N素限制。 （2）降雨量降低时，樟子松针叶的丙二醛（MDA）含量显著升高，针叶N含量降低，樟子松光合速率下降，同时，樟子松针叶的叶绿素含量大部分月份不受降雨量减少的影响，而且针叶脯氨酸和可溶性蛋白含量升高，超氧化物歧化酶（SOD）活性的升高，表明了樟子松对水分胁迫的生理生态适应机制。 （3）降雨量减少时樟子松林下植被总盖度显著降低，优势种由黄蒿和狗尾草演变为绿珠藜和黄蒿；降雨量增加时樟子松林下植被总盖度显著升高，优势种演变为艾蒿。降雨量减少和增加时物种多样性都显著降低，导致了生物多样性丧失。 （4）降雨量减少时樟子松和其林下植被的生长由于水分胁迫都受到了抑制，樟子松的高生长和粗生长速率减缓，林下植被的ANPP和地下部分生物量降低，进而导致樟子松人工林的地上部分C储量降低；樟子松的成长速率减缓和林下植被地上地下生物量的降低意味着生态系统凋落物量和死亡根系的减少，这直接导致了土壤有机碳含量的降低，即土壤有机碳储量的降低；综合降雨量减少导致的樟子松人工林的地上部分C储量降低和土壤有机碳储量的降低，我们的结果表明降雨量减少导致樟子松人工林C储量降低，同样的道理，降雨量增加导致樟子松人工林C储量升高。 （5）降雨量减少时，保护凋落物可以增加地表覆被，抑制地面水分蒸发，地表凋落物还能起到蓄水保水的作用，提高土壤水分有效性；降雨量增加时保护凋落物可以增加土壤养分（尤其是N）的输入，提高土壤养分的有效性。

Responses of Artemisia frigida Willd. (Compositae) and Leymus chinensis (Trin.) Tzvel. (Poaceae) to sheep saliva

Although studies show that grazing and browsing by herbivores have marked effects on host plants, the mechanisms remain unclear. The objective of this study is to determine the effects of sheep saliva on host plant growth. Sheep saliva was manually applied to clipped plants of two different life forms, a semi-shrub, Artemisia frigida Willd., and a herbaceous species, Leymus chinensis (Trin.) Tzevel. The results showed that sheep saliva significantly enhanced aboveground net primary productivity (ANPP) and the ratio of ANPP to belowground net primary productivity (BNPP) for both species. This indicated that sheep saliva promotes aboveground compensatory growth and allocation of photosynthate to aboveground for both plant species. Sheep saliva stimulated only tillering of L. chinensis. Regardless of saliva application, clipping significantly decreased BNPP and plant height, but significantly increased the number of branches or tillers for both plant species. The relative growth rates (RGRs) on both species were significantly greater after clipping with saliva compared with control and clipping without saliva treatments. In addition, RGR of the herbaceous species L. chinensis was faster than that of the semi-shrub A. frigida after application of saliva. (c) 2006 Elsevier Ltd. All rights reserved.

Experimental warming, not grazing, decreases rangeland quality on the Tibetan Plateau

We investigated experimental warming and simulated grazing ( clipping) effects on rangeland quality, as indicated by vegetation production and nutritive quality, in winter-grazed meadows and summer- grazed shrublands on the Tibetan Plateau, a rangeland system experiencing climatic and pastoral land use changes. Warming decreased total aboveground net primary productivity ( ANPP) by 40 g . m(-2) . yr(-1) at the meadow habitats and decreased palatable ANPP ( total ANPP minus non- palatable forb ANPP) by 10 g . m(-2) . yr(-1) at both habitats. The decreased production of the medicinal forb Gentiana straminea and the increased production of the non- palatable forb Stellera chamaejasme with warming also reduced rangeland quality. At the shrubland habitats, warming resulted in less digestible shrubs, whose foliage contains 25% digestible dry matter ( DDM), replacing more digestible graminoids, whose foliage contains 60% DDM. This shift from graminoids to shrubs not only results in lower- quality forage, but could also have important consequences for future domestic herd composition. Although warming extended the growing season in non- clipped plots, the reduced rangeland quality due to decreased vegetative production and nutritive quality will likely overwhelm the improved rangeland quality associated with an extended growing season.Grazing maintained or improved rangeland quality by increasing total ANPP by 20 - 40 g . m(-2) . yr(-1) with no effect on palatable ANPP. Grazing effects on forage nutritive quality, as measured by foliar nitrogen and carbon content and by shifts in plant group ANPP, resulted in improved forage quality. Grazing extended the growing season at both habitats, and it advanced the growing season at the meadows. Synergistic interactions between warming and grazing were present, such that grazing mediated the warming- induced declines in vegetation production and nutritive quality. Moreover, combined treatment effects were nonadditive, suggesting that we cannot predict the combined effect of global changes and human activities from single- factor studies.Our findings suggest that the rangelands on the Tibetan Plateau, and the pastoralists who depend on them, may be vulnerable to future climate changes. Grazing can mitigate the negative warming effects on rangeland quality. For example, grazing management may be an important tool to keep warming- induced shrub expansion in check. Moreover, flexible and opportunistic grazing management will be required in a warmer future.

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.

Inter-annual variability of precipitation constrains the production response of boreal Pinus sylvestris to nitrogen fertilization

© 2015 Published by Elsevier B.V.Tree growth resources and the efficiency of resource-use for biomass production determine the productivity of forest ecosystems. In nutrient-limited forests, nitrogen (N)-fertilization increases foliage [N], which may increase photosynthetic rates, leaf area index (L), and thus light interception (IC). The product of such changes is a higher gross primary production and higher net primary production (NPP). However, fertilization may also alter carbohydrate partitioning from below- to aboveground, increasing aboveground NPP (ANPP). We analyzed effects of long-term N-fertilization on NPP, and that of long-term carbon storing organs (NPPS) in a Pinus sylvestris forest on sandy soil, a wide-ranging forest type in the boreal region. We based our analyses on a combination of destructive harvesting, consecutive mensuration, and optical measurements of canopy openness. After eight-year fertilization with a total of 70gNm^-2, ANPP was 27±7% higher in the fertilized (F) relative to the reference (R) stand, but although L increased relative to its pre-fertilization values, IC was not greater than in R. On the seventh year after the treatment initiation, the increase of ANPP was matched by the decrease of belowground NPP (78 vs. 92gCm^-2yr^-1; ~17% of NPP) and, given the similarity of IC, suggests that the main effect of N-fertilization was changed carbon partitioning rather than increased canopy photosynthesis. Annual NPPS increased linearly with growing season temperature (T) in both treatments, with an upward shift of 70.2gCm^-2yr^-1 by fertilization, which also caused greater amount of unexplained variation (r²=0.53 in R, 0.21 in F). Residuals of the NPPS-T relationship of F were related to growing season precipitation (P, r²=0.48), indicating that T constrains productivity at this site regardless of fertility, while P is important in determining productivity where N-limitation is alleviated. We estimated that, in a growing season average T (11.5±1.0°C; 33-year-mean), NPPS response to N-fertilization will be nullified with P 31mm less than the mean (325±85mm), and would double with P 109mm greater than the mean. These results suggest that inter-annual variation in climate, particularly in P, may help explaining the reported large variability in growth responses to fertilization of pine stands on sandy soils. Furthermore, forest management of long-rotation systems, such as those of boreal and northern temperate forests, must consider the efficiency of fertilization in terms of wood production in the context of changes in climate predicted for the region.