利用方式对大针茅草原优势种糙隐子草空间异质性的影响

为探讨不同草地利用方式下糙隐子草种群空间分布的变化特点,以内蒙古自治区锡林郭勒盟毛登牧场大针茅草原优势物种糙隐子草(Cleistogenes squarrosa)为研究对象,通过野外试验,以地统计学为基础,采用半方差函数、分形维数及克里格插值法,分析了围封、放牧和刈割样地糙隐子草种群小尺度空间分布异质性。结果表明:糙隐子草种群密度分别为放牧(28.63株·m~(-2))围封(13.60株·m~(-2))刈割(3.85株·m~(-2))(P0.05),放牧显著增加糙隐子草种群密度(P0.05),而刈割显著降低糙隐子草种群密度(P0.05);通过半方差函数进行模型拟合,围封、放牧和刈割样地糙隐子草种群分布分别符合指数模型、球状模型和球状模型;空间分布格局分析表明,各样地糙隐子草种群的结构比分别为刈割(91.9%)放牧(75.3%)围封(71.1%),放牧与刈割样地的空间自相关性均较高,主要受结构性因素影响,而围封样地空间自相关处于中等程度,受随机性因素影响较大;结合分形维数、2D平面图及3D立体图分析,3块样地空间结构良好,空间分布简单,说明合理放牧及刈割不会对种群分布格局造成很大影响,但与围封样地相比,放牧增加了糙隐子草种群空间异质性,而刈割降低了种群异质性。     

草地利用方式对典型草原大针茅种群空间异质性的影响

以内蒙古锡林郭勒盟典型草原为研究对象,以野外实地调查的数据为基础,采用半方差函数和克里格插值法,分析了割草(M)、放牧(G)和围封(CK)样地中大针茅种群(Stipa grandis)小尺度空间分布异质性。结果表明:大针茅种群密度表现为割草样地最高,放牧样地次之,围封样地最低(P0.05);其中围封样地大针茅种群的结构比为95.3%,具有较强的空间自相关性,而刈割和放牧样地大针茅种群的结构比分别为70.2%和62.5%,属于中等程度的空间自相关性,大针茅种群的空间异质性表现为割草样地放牧样地围封样地;通过拟合半变异函数图,得到围封、放牧和割草样地大针茅种群空间分布最优模型分别为高斯模型、球状模型和球状模型;围封样地大针茅空间变异主要受结构因素的影响而趋于简单化分布格局,割草和放牧样地大针茅空间变异受到结构因素和随机因素的共同作用而表现为密度较大的斑块和密度较小的斑块镶嵌分布格局。     

刈割干扰下内蒙古草原两种丛生禾草繁殖策略的适应性调节

刈割是内蒙古草原的主要利用方式之一。当植物面对刈割干扰时, 可以通过繁殖策略的调节适应变化的环境。研究刈割干扰下植物繁殖策略的调节对于揭示植物的适应与进化机制具有重要意义。为此, 选择内蒙古典型草原的大针茅(Stipa grandis)和糙隐子草(Cleistogenes squarrosa), 在1979年围封的羊草(Leymus chinensis)样地(对照)和毗邻的长期刈割区对其繁殖策略进行了研究。结果显示: 1)长期刈割干扰下, 大针茅的种群生物量与对照区无显著差异, 由于其株丛生物量显著降低, 大针茅种群生物量的维持得益于种群密度的显著增加。糙隐子草的种群生物量比对照区显著提高, 由于其株丛生物量与对照区无显著差异, 糙隐子草种群生物量的增加也是因为密度的显著提高。因此, 密度调节是两种植物应对长期刈割干扰的重要的调节机制。2)长期刈割干扰下, 两种植物在无性繁殖和有性繁殖过程所采取的密度调节策略并不相同。大针茅种群生殖株丛的密度无显著变化, 非生殖株丛的密度显著提高, 导致其非生殖株丛的相对密度显著增加, 而生殖株丛的相对密度显著降低。因此刈割干扰下大针茅种群在密度调节上采取了保持有性繁殖株丛密度, 增加无性繁殖株丛密度的策略。糙隐子草种群非生殖株丛的密度显著增加, 但生殖株丛的密度增加幅度更大, 导致其非生殖株丛的相对密度显著降低, 而生殖株丛的相对密度显著提高。因此刈割干扰下糙隐子草种群在密度调节上采取了增加无性繁殖株丛密度的同时, 更大幅度地增加有性繁殖株丛密度的策略。3)长期刈割干扰下, 两种植物对生物量生殖分配的调节采取了不同的策略。大针茅株丛生物量向生殖枝分配的比例无显著变化, 通过降低生殖枝向穗的生物量分配比例, 降低了对有性生殖过程的生物量分配的投入; 而糙隐子草则通过增加株丛生物量向生殖枝分配的比例, 维持生殖枝向穗的生物量分配比例, 从而增加了对有性繁殖过程生物量分配的投入。4)长期刈割干扰下, 大针茅的株丛面积和株丛分蘖数与对照区无显著差异。糙隐子草株丛面积降低, 但是株丛分蘖数无显著变化。本实验说明, 植物种群有性繁殖和无性繁殖之间的密度调节和资源分配调节是草原植物应对刈割干扰下的重要调节机制, 不同的响应策略影响着植物种群的消长动态。     

不同恢复演替阶段糙隐子草种群的点格局分析

采用摄影定位法测定了羊草+大针茅草原不同恢复演替群落中糙隐子草种群的空间格局,并应用完全空间随机模型、泊松聚块模型和嵌套双聚块模型对其格局进行分析.结果表明:在严重退化的群落中,糙隐子草种群格局表现为嵌套双聚块结构,即在大聚块中分布着较高密度的小聚块;在恢复5年、8年和21年的群落中,则为以母体为中心的泊松聚块结构,即在糙隐子草种群空间格局的聚块中不存在较高密度的小聚块.这说明在严重退化的群落中正相互作用居主导,而在恢复演替群落中负相互作用居主导.糙隐子草种群在恢复演替过程中的格局变化主要是由于伴随放牧胁迫的消失,种群正相互作用(易化)向负相互作用(竞争)转化所致.     

白音华矿区草地群落主要物种组成及空间分布

以内蒙古白音华矿区周边草地为对象,研究了矿区草地植物群落的主要物种组成及其空间分布特征。结果表明: 草地群落共出现55种植物,优势种为大针茅、黄囊苔草和糙隐子草,常见种有羊草、冰草和知母等,该6个物种的累计相对重要值为79.6%,其密度分别为 26.6、204.7、105.4、107.1、68.2和55.1株·m^-2。对6个主要物种的种群密度通过半方差函数进行模型拟合, 其种群分布分别符合指数模型、指数模型、指数模型、球状模型、线性模型和高斯模型;对其空间分布格局进行分析,各种群的结构比分别为59.2%、97.2%、89.1%、94.5%、62.6%和72.1%,表明黄囊苔草、糙隐子草和羊草种群的空间自相关性程度均较高, 主要受结构性因素影响, 而大针茅、冰草和知母种群主要受随机性因素影响。对分形维数进行分析发现, 大针茅、黄囊苔草、糙隐子草和冰草种群分布格局较简单,空间依赖性较强,而羊草和知母种群分布格局较复杂,空间依赖性较弱,结合2D及3D图看, 大针茅和知母呈现出梯度扩散,而黄囊苔草、糙隐子草、羊草和冰草则主要呈现斑块化分布,表明矿区草地群落主要物种的空间分布与开矿无显著关联性。     

典型草原大针茅种群空间格局及对长期过度放牧的响应

种群空间格局是生态学研究的基本问题之一。典型草原带由于过度放牧退化严重, 原生群落罕见, 探讨原生群落的种群空间格局具有重要生态学意义。大针茅(Stipa grandis)草原是典型草原区广泛分布的主要群落类型, 1979年围封的大针茅样地, 是目前保存完整的大针茅草原原生群落。本文选择大针茅草原原生群落和长期过度放牧群落, 应用O-Ring函数结合不同零假设模型分析了大针茅种群的空间格局。结果表明: 在原生群落中大针茅种群在小尺度范围内呈均匀分布, 而在长期过度放牧群落中则表现为聚集分布。这说明在大针茅草原原生群落中竞争是主要的相互作用, 而在长期过度放牧群落中正相互作用居主导, 验证了胁迫梯度假说; 同时证明长期过度放牧改变了种群空间格局。     

内蒙古大针茅草原常见植物在不同土地利用方式下的固碳相关属性

为了探究植物与固碳相关属性在不同功能群、器官和物种间的差异, 以及这些属性对不同土地利用方式的响应, 2012年8月对内蒙古大针茅草原的4个样地(长期无干扰样地、长期自由放牧样地、4年围封样地和4年围封割草样地)进行了群落调查, 并采集样地中的常见植物, 测定了与植物固碳相关的属性, 包括全碳含量、全氮含量、碳氮比、纤维素含量、木质素含量和酸性洗涤纤维含量等。以常见植物为对象, 在功能群水平研究了各土地利用方式下全碳含量、全氮含量和碳氮比的差异; 针对各样地的共有种——糙隐子草(Cleistogenes squarrosa)、大针茅(Stipa grandis)、冷蒿(Artemisia frigida)、羊草(Leymus chinensis)和猪毛菜(Salsola collina), 从物种和器官水平分析了不同土地利用方式下植物的固碳相关属性。结果表明: 大针茅草原植物不同功能群、物种和器官间的固碳相关属性存在极显著差异, 不同土地利用方式下大针茅草原不同功能型、物种和器官的固碳相关属性有显著差异。与其他利用方式相比, 4年围封割草对植物与固碳相关属性的影响最为明显, 功能群、器官和物种水平的植物氮含量均有下降。糙隐子草和猪毛菜的这些属性对长期自由放牧的响应敏感, 且方向相反。     

Traits related to carbon sequestration of common plant species in a Stipa grandis steppe in Nei Mongol under different land-uses

为了探究植物与固碳相关属性在不同功能群、器官和物种间的差异, 以及这些属性对不同土地利用方式的响应, 2012年8月对内蒙古大针茅草原的4个样地(长期无干扰样地、长期自由放牧样地、4年围封样地和4年围封割草样地)进行了群落调查, 并采集样地中的常见植物, 测定了与植物固碳相关的属性, 包括全碳含量、全氮含量、碳氮比、纤维素含量、木质素含量和酸性洗涤纤维含量等。以常见植物为对象, 在功能群水平研究了各土地利用方式下全碳含量、全氮含量和碳氮比的差异; 针对各样地的共有种——糙隐子草(Cleistogenes squarrosa)、大针茅(Stipa grandis)、冷蒿(Artemisia frigida)、羊草(Leymus chinensis)和猪毛菜(Salsola collina), 从物种和器官水平分析了不同土地利用方式下植物的固碳相关属性。结果表明: 大针茅草原植物不同功能群、物种和器官间的固碳相关属性存在极显著差异, 不同土地利用方式下大针茅草原不同功能型、物种和器官的固碳相关属性有显著差异。与其他利用方式相比, 4年围封割草对植物与固碳相关属性的影响最为明显, 功能群、器官和物种水平的植物氮含量均有下降。糙隐子草和猪毛菜的这些属性对长期自由放牧的响应敏感, 且方向相反。     

放牧对短花针茅荒漠草原建群种与优势种空间分布关系的影响

采用多种分析方法探讨放牧对短花针茅荒漠草原建群种与优势种无芒隐子草、碱韭的种群空间分布关系的影响.结果表明: 短花针茅+无芒隐子草+碱韭群落的景观特征主要由草原的土壤理化性质和种群的固有属性决定.自由放牧导致的空间分布的变异程度为碱韭>无芒隐子草>短花针茅.自由放牧对短花针茅和碱韭密度的影响显著,但对无芒隐子草影响不大.随着无芒隐子草密度的增加,自由放牧区短花针茅密度呈现增大趋势,而围封区短花针茅种群密度呈现先增大后减小的趋势;自由放牧区和围封区内短花针茅密度均随碱韭密度的增加而减小.表明植物种群种间关系存在密度效应,在受到外界干扰的情况下,密度效应可能消失;因植物种对或干扰条件不同,植物种间关系可以有多种表现形式.     

氮素添加和功能群去除对糙隐子草和大针茅根系特征的影响

由于草地不合理利用,中国北方草原严重退化,导致生态系统结构性破坏甚至功能性紊乱。随着草原退化,中能值功能群植物糙隐子草会取代高能值功能群植物大针茅成为群落中优势度最大的植物。为了解大针茅和糙隐子草在氮素添加以及去除功能群的情况下根系特征的变化趋势,选取锡林郭勒盟典型草原中的退化草原群落,分别开展两项实验,实验1:氮素添加梯度实验(0、10.5、17.5、28 gN/m~2),实验2:同时进行功能群的去除和氮素的添加,功能群去除实验(将植物分为高能值植物功能群、中能值植物功能群、低能值植物功能群3个功能群;每个处理分别剪除另外两个功能群,留取单一功能群),并同时进行氮素添加实验(0、17.5 gN/m~2)。在进行两年的实验处理后,通过Delta-T SCAN根系分析仪测量大针茅和糙隐子草根系的长度、直径、面积、体积指标。分析植物根系对氮素添加的响应,以及功能群去除是否改变这两种植物对氮素添加的响应格局。实验1研究结果表明:在受到其他功能群的竞争压力下,大针茅根系长度、面积、体积均在高氮素添加(28 gN/m~2)情况下显著增加,糙隐子草根系直径和体积在中氮素添加(17.5 gN/m~2)的情况下显著高于其他3个处理,退化样地中土壤氮素的增加,促使大针茅根系主要通过增加根系长度扩大在土壤中的空间分布,而促使糙隐子草主要通过增粗生长来扩大土壤分布空间。实验2研究结果表明:功能群的去除,中氮素添加对根系的影响,只有糙隐子草的根系直径显著增加。综合来看,功能群去除实验对大针茅的根系长度和面积有影响,对糙隐子草的根系长度、直径和面积有影响。功能群去除处理与氮素添加的交互作用对大针茅根系没有影响,对糙隐子草的根系直径和体积有影响。     

Effects of grazing on photosynthetic characteristics of major steppe species in the Xilin River Basin,Inner Mongolia,China

Species composition and photosynthetic characteristics of dominant species of ungrazed plot (UG), overgrazed plot (OG), and restored grazed plot (RG) were determined in the Xilin River Basin, Inner Mongolia, China. Both heavily grazing and restoration significantly affected the composition of different species and life forms. Leymus chinensis, Stipa grandis, and Cleistogenes polyphylla, three dominant perennial grasses in UG plot, contributed 58.9 % aboveground biomass to that of whole community, and showed higher net photosynthetic rate (P _N), transpiration rate (E), and intrinsic water-use efficiency (WUE). In OG plot, relative biomass of L. chinensis and S. grandis significantly decreased, while relative biomass of three shrubs/sub-shrubs, Caragana microphylla, Artemisia frigida, and Kochia prostrata, obviously increased. Heavy grazing significantly decreased P _N, E, and WUE of L. chinensis and S. grandis, while shrubs/sub-shrubs showed significantly higher photosynthetic activity and WUE than the grasses. After 18-year restoration, photosynthetic activities of L. chinensis and S. grandis were significantly higher than those in the OG plot. The proportion of L. chinensis, S. grandis, and C. microphylla significantly increased, and relative biomass of C. polyphylla, A. frigida, and K. prostrata markedly declined in RG plot. We found close relationships between physiological properties of species and their competitive advantage in different land use types. Higher photosynthetic capability means more contribution to total biomass. The variations in physiological characteristics of plants could partly explain the changes in species composition during degrading and restoring processes of Inner Mongolia typical steppes.     

Effect of long-term grazing on soil organic carbon content in semiarid steppes in Inner Mongolia

To clarify the response of soil organic carbon (SOC) content to season-long grazing in the semiarid typical steppes of Inner Mongolia, we examined the aboveground biomass and SOC in both grazing (G-site) and no grazing (NG-site) sites in two typical steppes dominated by Leymus chinensis and Stipa grandis, as well as one seriously degraded L. chinensis grassland dominated by Artemisia frigida. The NG-sites had been fenced for 20 years in L. chinensis and S. grandis grasslands and for 10 years in A. frigida grassland. Aboveground biomass at G-sites was 21–35% of that at NG-sites in L. chinensis and S. grandis grasslands. The SOC, however, showed no significant difference between G-site and NG-site in both grasslands. In the NG-sites, aboveground biomass was significantly lower in A. frigida grassland than in the other two grasslands. The SOC in A. frigida grassland was about 70% of that in L. chinensis grassland. In A. frigida grassland, aboveground biomass in the G-site was 68–82% of that in the NG-site, whereas SOC was significantly lower in the G-site than in the NG-site. Grazing elevated the surface soil pH in L. chinensis and A. frigida communities. A spatial heterogeneity in SOC and pH in the topsoil was not detected the G-site within the minimal sampling distance of 10 m. The results suggested that compensatory growth may account for the relative stability of SOC in G-sites in typical steppes. The SOC was sensitive to heavy grazing and difficult to recover after a significant decline caused by overgrazing in semiarid steppes.     

Asymmetric effects of grazing intensity on macroelements and microelements in grassland soil and plants in Inner Mongolia Grazing alters nutrient dynamics of grasslands

Grazing is a traditional grassland management technique and greatly alters ecosystem nutrient cycling. The effects of grazing intensity on the nutrient dynamics of soil and plants in grassland ecosystems remain uncertain, especially among microelements. A 2‐year field grazing experiment was conducted in a typical grassland with four grazing intensities (ungrazed control, light, moderate, and heavy grazing) in Inner Mongolia, China. Nutrient concentration was assessed in soil and three dominant plant species (Stipa krylovii, Leymus chinensis, and Cleistogenes squarrosa). Assessed quantities included four macroelements (carbon (C), nitrogen (N), phosphorus (P), and magnesium (Mg)) and four microelements (copper (Cu), iron (Fe), manganese (Mn), and zinc (Zn)). Soil total C, total N, total P, available N, and available P concentrations significantly increased with grazing intensity but soil Mg, Cu, Fe, Mn, Zn concentrations had no significant response. Plant C concentration decreased but plant N, P, Mg, Cu, Fe, Mn, and Zn concentrations significantly increased with grazing intensity. In soil, macroelement dynamics (i.e., C, N, and P) exhibited higher sensitivity with grazing intensity, conversely in plants, microelements were more sensitive. This result indicates macroelements and microelements in soil and plants had asymmetric responses with grazing intensity. The slopes of nutrient linear regression in C. squarrosa were higher than that of S. krylovii and L. chinensis, indicating that C. squarrosa had higher nutrient acquisition capacity and responded more rapidly to heavy grazing. These findings indicate that short‐term heavy grazing accelerates nutrient cycling of the soil–plant system in grassland ecosystems, elucidate the multiple nutrient dynamics of soil and plants with grazing intensity, and emphasize the important function of microelements in plant adaptation in grazing management.     

Above- and belowground biomass allocation and its regulation by plant density in six common grassland species in China

Above- and belowground biomass allocation is an essential plant functional trait that reflects plant survival strategies and affects belowground carbon pool estimation in grasslands. However, due to the difficulty of distinguishing living and dead roots, estimation of biomass allocation from field-based studies currently show large uncertainties. In addition, the dependence of biomass allocation on plant species, functional type as well as plant density remains poorly addressed. Here, we conducted greenhouse manipulation experiments to study above- and belowground biomass allocation and its density regulation for six common grassland species with different functional types (i.e., C₃ vs C₄; annuals vs perennials) from temperate China. To explore the density regulation on the biomass allocation, we used five density levels: 25, 100, 225, 400, and 625 plant m^?2. We found that mean root to shoot ratio (R/S) values ranged from 0.04 to 0.92 across the six species, much lower than those obtained in previous field studies. We also found much lower R/S values in annuals than in perennials (C. glaucum and S. viridis vs C. squarrosa, L. chinensis, M. sativa and S. grandis) and in C₄ plants than in C₃ plants (C. squarrosa vs L. chinensis, M. sativa and S. grandis). In addition to S. grandis, plant density had significant effects on the shoot and root biomass fraction and R/S for the other five species. Plant density also affected the allometric relationships between above- and belowground biomass significantly. Our results suggest that R/S values obtained from field investigations may be severely overestimated and that R/S values vary largely across species with different functional types. Our findings provide novel insights into approximating the difficult-to-measure belowground living biomass in grasslands, and highlight that species composition and intraspecific competition will regulate belowground carbon estimation.

     

Arbuscular mycorrhizal fungi alter plant community composition along a grazing gradient in Inner Mongolia Steppe

Arbuscular mycorrhizal fungi (AMF) have a significant influence on plant productivity and diversity in non-grazing grassland. However, the interactive effects between grazing intensity and AMF on plant community composition in natural grassland communities are not well known. We conducted a field experiment that manipulated AMF colonization and grazing intensity to study the impact of AMF suppression on plant community composition and nutrient status over 2 years (2015–2016) with contrasting rainfall levels. We found that AMF root colonization was significantly reduced by the application of the fungicide benomyl as a soil drench. Grazing intensity regulated plant community composition and aboveground biomass mainly by reducing the growth of Leymus chinensis over 2 years. AMF suppression increased the growth of Chenopodium glaucum, but it did not alter other plant species across all grazing intensities. The effects of AMF suppression on plant community composition changed along a grazing gradient considerably between years: AMF suppression increased the biomass of C. glaucum across all grazing intensities in 2015, but slightly increased it in 2016. Interactions between AMF suppression and grazing intensity altered the phosphorus concentration of Stipa grandis and Cleistogenes squarrosa in 2015 but not in 2016. AMF suppression decreased the shoot phosphorus content of L. chinensis but increased that of C. glaucum across all grazing intensities. Our results indicate that grazing intensity substantially alters aboveground community biomass and affects growth of dominant species; AMF by itself have limited effects on plant communities along a grazing gradient in typical steppe.     

羊草与大针茅根系构型对水分梯度响应的比较研究

羊草(Leymus chinensis)与大针茅(Stipa grandis)是内蒙古锡林郭勒典型草原的两大建群种,也是内蒙古草原的重要优良牧草。选取锡林郭勒草原以大针茅和羊草为优势种的围封草场为研究样地,通过原状土柱移栽,进行了两年的水分梯度控制实验(150、300、450、600 mm),分别模拟当地年降雨量由干旱年到是湿润年的变化情况,分析比较羊草种群和大针茅种群地上高度和根系构型对水分梯度的响应情况。结果表明:相比大针茅种群,羊草种群对水分梯度的响应更敏感,随着水分梯度的增加,羊草地上高度和根系直径显著增加,根系长度和根系深度显著减少;而大针茅的地上高度和根系特征各项指标均没有显出与水分梯度的相关性,显然大针茅比羊草更能适应干旱生境。在干旱条件(模拟年降雨量150 mm处理)下,大针茅种群地上高度达到峰值,生长状况良好;羊草种群则采用地上个体小型化,地下主根变细,分叉,向土壤深层扩展的生长策略。因此,在气候变化背景下,干旱化的气候将导致大针茅种群在群落中的优势地位逐渐增加,反之羊草种群则会随着气候湿润化而占据更大的优势。     

Response of dominant grassland species in the temperate steppe of Inner Mongolia to different land uses at leaf and ecosystem levels

In order to study the responses of dominant species to different land uses in the semiarid temperate grassland of Inner Mongolia, we tested the physiological responses of Stipa grandis, Leymus chinensis, and Artemisia frigida to mowing, grazing exclusion, and grazing land uses at the leaf and ecosystem levels. The grazing-exclusion and mowing sites released CO₂, but the grazing site was a net carbon sink. L. chinensis and S. grandis contributed more to the ecosystem CO₂ exchange than A. frigida. At the grazing-exclusion and mowing sites, Leymus chinensis and Stipa grandis both exhibited a higher light-saturation point and higher maximum photosynthetic rate than that at the grazing site, which increased photosynthesis and growth compared to those at the grazing site. In contrast, A. frigida possessed a higher nitrogen content than the other species, and more of the light energy used for photosynthesis, particularly at the grazing site.     

Short-term regrowth responses of four steppe grassland species to grazing intensity, water and nitrogen in Inner Mongolia

Inner Mongolia steppe grasslands are widely used for livestock farming and the regrowth ability of grassland species is therefore strongly influenced not only by water and nutrient availability but also quite heavily by grazing. However, little is known on how grazing, water and nitrogen interactively affect the dominant C3 species (Leymus chinensis, Stipa grandis, Agropyron cristatum) and the C4 species (Cleistogenes squarrosa). Therefore in the 2007 and 2008 growing seasons, a field experiment was carried out to test the hypothesis that under different grazing intensities the dominant species show different short-term regrowth response to simultaneous variation in the availability of water and nitrogen. Single factor and interaction effects of the addition of water (rainfed vs. simulated wet-year) and nitrogen (0 or 25 kg N ha^?1) were analysed along a gradient of four grazing intensities (ungrazed, lightly, moderately and heavily grazed) after one month of grazing exclusion. Water and nitrogen addition affected short-term regrowth of all species in a similar way whereas species responded differently to grazing. Simulated wet-year water availability consistently resulted in higher standing biomass, relative growth rate and cellulase digestible organic matter yield. Supplementary nitrogen promoted standing biomass and crude protein concentration. The nutritive value of all species’ standing biomass showed a similar increase with more intensive grazing. However, heavy grazing led to a clear shift in the relative biomass of the species, i.e. mainly a promotion of the C4 grass, C. squarrosa. In contrast to our hypothesis, there were no differences among species in their response to water or nitrogen addition, whereas, heavy grazing induced the expected species-specific response. Our results suggest that heavy grazing rather than nitrogen or water determine short-term shifts in species composition of the investigated steppe ecosystem. Furthermore, differences in the species-specific growth response to grazing may increase the proportion of the C4 grass C. squarrosa in steppe communities, whereas higher availability of nitrogen and water may lead to higher forage biomass and nutritive value of all investigated species but in short-term cannot compensate for the grazing induced changes in species composition.     

Effects of grazing management system on plant community structure and functioning in a semiarid steppe: scaling from species to community

Under the aim of searching for a more sustainable grazing management system, a mixed management system (grazing and haymaking alternate annually) was proposed and tested against traditional management system (used consistently either for grazing or haymaking) in the semiarid grassland of Inner Mongolia with a field manipulation experiment. The responses of aboveground biomass to the two grazing management systems were examined across different levels of organization (i.e., species, plant functional group, and community) and in five consecutive years from 2005 to 2009. The effects of the two systems on seed production potential of four dominant species (Leymus chinensis, Stipa grandis, Agropyron cristatum, Cleistogenes squarrosa) were also investigated. Our results demonstrate that, in the traditional system, aboveground biomass production across all the levels of organization was reduced by grazing. In mixed system, however, no significantly negative relationship between the biomass response and stocking rate was detected at all organization levels. Precipitation fluctuation had strong influence on biomass responses, and compared to the traditional system the slope of the biomass-precipitation relationship tends to be higher in the mixed system. This effect might be attributed to the more positive response of L. chinensis and A. cristatum to increase in precipitation. In the traditional system, both the ratio and the density of reproductive tillers of the grazing subplots were significantly reduced compared to the haymaking or ungrazed control plots. In the mixed system, there was no significant difference between the haymaking subplots and the ungrazed control plots, regardless of the grazing pressures imposed on the haymaking subplots in the previous growing season. Our findings suggest that the mixed system mitigates the sheep grazing-induced species shift and it tends to be more responsive to increasing precipitation as compared to the traditional system. Therefore, replacement of the traditional grazing strategy with the mixed system could provide an important contribution to sustainable land-use of the Inner Mongolia grasslands.