高寒矮嵩草草甸两种主要植物耐牧性的比较

通过野外控制实验,研究了刈割(留茬3 cm、留茬1 cm及不刈割)、施肥(施肥、不施肥)和浇水(浇水、不浇水)处理对高寒草甸矮嵩草(Kobresia humilis)和垂穗披碱草(Elymus nutans)补偿高度、株高相对生长率、比叶面积、叶片净光合速率和地上总生物量的影响.结果表明:2物种的株高和地上总生物量在刈割后均为低补偿响应,但其株高相对生长率显著提高,并均随年份而增加;垂穗披碱草比叶面积、叶片净光合速率和地上总生物量对刈割损伤更加敏感;尽管施肥能显著提高2物种上述各项指标,但在不同处理条件下矮嵩草的耐牧性指数均小于垂穗披碱草;浇水的作用不显著.说明2物种的耐牧性依赖于土壤养分资源获得性,矮嵩草的耐牧性强于垂穗披碱草.     

刈割、施肥和浇水对垂穗披碱草补偿生长的影响

植物的补偿生长特性受放牧强度和生境资源获得性的影响。通过为期2年的野外控制实验,研究了刈割高度(留茬1cm、3cm及不刈割)、施肥(施、不施)和浇水(浇、不浇)处理对垂穗披碱草(Elymus natans)补偿生长的影响,并结合对各处理分株密度、比叶面积、净光合速率和相对生长率的变化研究,探讨了其补偿生长机制。结果表明:刈割后垂穗披碱草分株种群密度显著增加,补偿生长高度显著降低,比叶面积和相对生长率随刈割强度增加而呈上升趋势,叶片净光合速率变化不显著;施肥能显著增加垂穗披碱草的补偿生长高度、比叶面积、叶片净光合速率和相对生长率;浇水处理以及刈割、浇水、施肥处理之间的交互作用均不显著。可见,在刈割条件下,垂穗披碱草具有一定的密度补偿机制,但由于刈割抑制补偿性高生长,导致分株高度出现低补偿。因此,即使刈割后比叶面积和株高相对生长率显著增加,也不一定必然引起株高的超补偿;但施肥可显著提高垂穗披碱草的补偿能力,增加耐牧性,证实了改进后的限制资源模型的预测。     

克隆植物矮嵩草在刈割条件下的等级反应研究

对刈割处理下高寒矮嵩草草甸建群种矮嵩草（Kobresia humilis）构件等级中基株、分株和分蘖3个层次构件单元的数量、生物量及其变异系数的变化进行了研究。结果表明．矮嵩草株丛刈割部分和未刈割部分被测性状在处理间的差异显著性以及作构件等级间变异系数的大小均为分蘖层次〉分株层次〉基株层次。刈割后相同时期内,分蘖层次对相同处理的反应发叶生得最快,基株则最为迟缓。这说明在刈割后的相同时期内,构件等级中较外层次（分蘖层次）会发生更大的表型变异．对相同强度的刈割扰动发生反应也最快。由此证实矮嵩草在刈割条件下也同样会发生等级性反应。     

高寒草甸对刈割、施肥和浇水发生响应的最优植物性状集和功能型

基于植物性状和功能型的特征变化对于研究植被动态和生态系统功能变化具有重要意义。通过在高寒矮嵩草(Kobresia humilis)草甸为期5年(2007-2011年)的刈割(不刈割、留茬3 cm、留茬1 cm)、施肥(施肥、不施肥)和浇水(浇水、不浇水)控制实验, 采用递归算法(recursive algorithm)和多元回归分析筛选对模拟放牧发生响应的最优植物性状集和响应功能型, 以及影响群落生产力变化的作用功能型。研究结果显示: (1)在不施肥不浇水、仅施肥、仅浇水和既施肥又浇水4种条件下的最优植物性状集不同, 它们分别是叶缘形状-株高-叶干质量-比叶面积、生活周期-株高-叶干质量-比叶面积、生活周期-叶片叶绿素含量-叶表面结构-株高-叶干质量-比叶面积和繁殖结构-叶缘-株高。其中, 株高、叶干质量和比叶面积是对刈割和土壤资源变化更为敏感的植物性状。(2)在这4种处理条件下, 共获得14个最优响应功能型和4个作用功能型。作用功能型对群落生产力变异的解释能力在50.3%-86.4%之间。(3)最优响应功能型和作用功能型分别占功能型总数的70%和20%。作用功能型占最优响应功能型的28.5%, 两者间仅存在部分重叠。上述结果说明, 植物功能性状和功能型变化能够准确地反映植被的放牧响应和生态系统功能变化, 但是不同资源条件下群落的最优响应性状集和功能型不同。作用功能型是同时反映植被放牧响应和生态系统功能变化的最优功能型。     

Assembly patterns of plant functional traits in alpine meadow under disturbances by mowing and fertilization

针对目前关于植物群落功能性状构建模式及其驱动因素存在的争议, 研究了33个物种10个功能性状的构建模式。研究结果显示: (1)在刈割-施肥复合梯度上, 这些功能性状主要表现为随机构建模式, 发生随机性和确定性构建的样方比例分别为82.7%和17.3%; (2)在10个功能性状中, 生长型、生活周期、单株地上干质量、叶面积和叶干质量5个功能性状为随机构建模式, 不受试验处理和群落特征(地上净初级生产力、刈割生物量损失、群落高度)变化的影响。植物倾斜度、繁殖方式、固氮性、株高和比叶面积5个功能性状的构建模式与试验处理或群落特征变化有关, 表现出趋同或趋异构建响应。其中, 植物倾斜度和比叶面积的构建模式仅受群落特征影响, 而固氮性、繁殖方式和株高3个性状的构建模式同时还受刈割或施肥处理的影响, 其构建模式因功能性状而异; (3)群落的生物量损失和地上净初级生产力是解释植物功能性状构建模式变化较理想的群落特征; (4)刈割和施肥处理对株高的构建模式具有相反效应, 而刈割和施肥的交互作用对其无显著影响。上述结果说明该研究群落植物功能性状的构建存在不同模式, 以随机构建模式为主, 确定性构建模式居次要地位。确定性构建模式与试验处理和群落特征变化有关, 而且是性状依赖的。相反选择力对趋同和趋异构建模式的平衡效应能引起功能性状发生随机构建。     

光照，施肥及刈割对垂穗披碱草生长的影响

以甘南高寒草甸常见牧草垂穗披碱草（Elymus nutans）为研究对象，比较不同光照、肥力条件下，垂穗披碱草对刈割的生长和补偿反应。研究发现，光照和肥力存在交互作用。施肥、中度光照下，未刈割单株垂穗披碱草地上生物量最大，为3．239g。施肥、中度光照下，比较刈割植株地上生物量补偿发现：晚期刈割单株地上生物量的补偿大于早期刈割；刈割强度无显著影响；刈割两次单株地上生物量的补偿最大。施肥、中度光照、晚期、轻度、一次刈割时，单株垂穗披碱草地上生物量累积最大，为4．500g。全光照、未施肥条件下，刈割对单株地上生物量的补偿无影响。就补偿机制而言，植株地上生物量的增加以地下生物量为代价。     

施肥和刈割对冷地早熟禾补偿生长的影响

以青藏高原东部高寒草甸常见牧草冷地早熟禾(Poa crymophila)为研究对象,比较研究了肥力和不同刈割处理对植物生长和生物量补偿的影响。实验采用随机设计,肥力分为施肥和不施肥,刈割处理为刈割时间、刈割频度和刈割强度3因子组合。结果表明:不刈割情况下,施肥显著促进冷地早熟禾的生长,而在刈割情况下,施肥对植物的生长没有明显的促进作用甚至有负作用。不施肥,刈割可促进冷地早熟禾地上生物量的增加并发生超补偿,但补偿程度因刈割强度、刈割频度和刈割时间而异;施肥时,刈割没有或很少促进植株发生超补偿。超补偿发生时,资源贫瘠的条件可能是必需的。冷地早熟禾发生超补偿可能是以损耗地下部分的资源为代价的,刈割刺激分蘖的增加对超补偿的发生也有一定的贡献。     

Effects of fertilization and clipping on species diversity,productivity and their relationship in subalpine meadow

施肥和刈割分别对植物群落物种多样性和生产力有重要的影响。采用不同施肥水平和刈割频度, 研究了施肥和刈割对亚高山草甸植物群落物种多样性与生产力及其关系的影响。结果显示, 5年的施肥和刈割处理对群落地上生物量均有显著影响, 但对物种多样性影响不显著; 物种多样性与生产力之间的关系因施肥和刈割次数的不同而不同, 有负相关、二次函数关系和不相关等几种类型, 多数为不相关。此结果表明施肥和刈割并不总是一致性地影响群落物种多样性与生产力。因此, 不同施肥和刈割处理下的亚高山草甸植物群落生产力与物种多样性之间并没有确定的关系。     

青藏高原高寒草甸植物群落结构和功能对氮、磷、钾添加的短期响应

对中国科学院海北高寒草甸生态系统定位站的矮嵩草草甸水肥样地进行了氮、磷、钾及其组合的施肥处理,研究了施肥对植物群落结构和功能的影响。结果表明:(1)施肥使矮嵩草草甸植物群落物种丰富度减少,不同施肥处理下物种丰富度大小分别为:对照钾磷氮氮磷磷钾氮钾氮磷钾。(2)在氮磷配合施肥处理下,矮嵩草草甸植物群落Shannon-Wiener指数显著高于对照,而其它施肥处理对Shannon-Wiener指数影响不显著。(3)在同一施肥处理下,禾草类和莎草类的重要值明显高于豆科和杂类草功能群,不同施肥处理使禾草、莎草、豆科植物的重要值增加,而杂类草重要值减少。(4)与对照相比,不同施肥(除钾外)处理可不同程度的增加植物群落的高度。(5)除钾、磷钾养分添加对矮嵩草草甸地上生物量的影响与对照差异不显著外,其它养分及其组合添加都极显著增加了群落地上生物量,且大小顺序依次为氮磷氮磷钾磷氮钾氮磷钾钾对照。(6)施用不同种类的肥料后,矮嵩草草甸各功能群地上生物量的比例变化明显,禾草和莎草的比例均增加,杂类草的比例减少,而豆科植物无规律性。(7)熵值法综合评价短期施肥处理对矮嵩草草甸群落的影响表明,氮磷、氮磷钾配合施肥是青藏高原高寒草甸最佳施肥选择。     

不同放牧强度下矮嵩草（Kobresia humilis）无性系分株种群的动调

以矮嵩草无性系分株和分蘖分别作为其种群的基本单元,对不同放牧强度下种群的动态与调节进行了研究。结果表明：随着放牧强度的增加,每分株的分蘖数、叶片数及分株个体地上生物量均增加．分蘖死亡率和叶片死亡率在各处理间差异不显著。分蘖死亡率的高峰出现在生长季末,叶片死亡率在生长初期和末期较高,而且都不属于密度制约性的死亡。矮嵩草多重种群结构水平的数量调节是由最外层次（叶片层次）的数量变化引起的,进而影响到较内层次上结构单元的大小和数量。     

Compensatory growth responses to clipping defoliation in Leymus chinensis (Poaceae) under nutrient addition and water deficiency conditions

Compensatory growth responses of Leymus chinensis, a dominant species in Inner Mongolia steppe, to clipping defoliation were evaluated in a pot-cultivated experiment under different nutrient (N and P) and water availability conditions. Leymus chinensis exhibited over-compensatory growth at the light and moderate clipping intensities (20% and 40% aerial mass removed) with a greater accumulated aboveground biomass, higher relative growth rate (RGR), more rhizomatic tillers and a stimulation of compensatory photosynthesis to the remnant leaves as compared with those of the unclipped plants. Intense clipping (80% aerial mass removed), which removed most of the aboveground tissues, greatly reduced the growth of aboveground biomass in comparison with that of the unclipped plants. Nitrogen addition only slightly improved the biomass production and RGR in light and moderately clipped plants, and it did not allow plants in the intense clipping condition to over-compensate. Phosphorus addition had no obvious influences on the growth and physiological responses to clipping defoliation. These results indicated that nutrient addition could not compensate for the negative effects of severe clipping on the defoliated grass. On the other hand, there were no distinct positive responses under water deficiency condition for L. chinensis at all clipping intensities with a significant reduction of aboveground and belowground biomass, lower RGR, fewer rhizomatic tillers, and a lower net photosynthetic rate than other wet treatments. Additionally, the chlorophyll contents of remnant leaves gradually increased with the increase of clipping intensities in each treatment. In conclusion, although L. chinensis could compensate for tissues removal by some morphological and physiological responses, intense clipping and drought can result in a significant decrease of biomass and growth rate, even under enriched nutrition conditions.     

Clonal integration improves compensatory growth in heavily grazed ramet populations of two inland-dune grasses

Herbaceous species possess several mechanisms to compensate for tissue loss. For clonal herbaceous species, clonal integration may be an additional mechanism. This may especially hold true when tissue loss is very high, because other compensatory mechanisms may be insufficient. On inland dunes in northern China, we subjected Bromus ircutensis and Psammochloa villosa ramets within 0.5 m×0.5 m plots to three clipping treatments, i.e., no clipping, moderate (50% shoot removal) and heavy clipping (90% shoot removal), and kept rhizomes at the plot edges connected or disconnected. Moderate clipping did not reduce ramet, leaf or biomass density of either species. Under moderate clipping, rhizome connection significantly improved the performance of Psammochloa, but not that of Bromus. Heavy clipping reduced ramet, leaf and biomass density in the disconnected plots of both species, but such negative effects were negated or greatly ameliorated when the rhizomes were connected. Therefore, clonal integration contributed greatly to the compensatory growth of both species. The results suggest that clonal integration is an additional compensatory mechanism for clonal plants and may be important for their long-term persistence in the heavily grazed regions in northern China.     

叶片损害强度与基质营养水平对苦草补偿性生长的影响

通过室外双因子控制试验,研究了叶片损害强度与基质营养水平对沉水植物苦草补偿性生长的影响.结果表明：叶片损害与基质类型对苦草的累积生物量、生物量的分配（根叶比）及分株数均有显著影响.与未受损植物相比，强受损使苦草的累积生物量及无性分株数显著降低，根叶比明显升高，而弱受损与未受损处理之间没有明显差别.与营养贫瘠的岸泥相比，生长在肥力较高湖泥中的苦草的累积生物量较高，而根叶比与分株数较低；叶片损害与基质类型对苦草的相对生长率也有显著影响，受损使苦草的生长率明显提高，而两受损处理之间无显著差别.未受损与强受损时,湖泥中苦草的生长速率明显高于岸泥，弱受损时则相近.并初步探讨了水生植物受损后的补偿生长机理.     

Interspecific variations in responses of <Emphasis Type="Italic">Festuca rubra</Emphasis> and <Emphasis Type="Italic">Trifolium pratense</Emphasis> to a severe clipping under environmental changes

Understanding and predicting possible responses of grassland species to global change is of important meaning for adapting grassland management to a changed and changing environment. A laboratory clipping experiment was conducted to examine the interspecific responses in an ecological context of competition and environmental changes. Festuca rubra and Trifolium pratense, either in monoculture or two- and three-species mixtures, were grown in three environmental combinations (ambient and increased temperature, repetitive N supply, and simulated acid rain), respectively. After a growth time of three months, plants were clipped at the height of 1.0 cm above soil surface. Plant height and aboveground biomass prior clipping, and survival rate and regrowth (height and biomass) after clipping were analyzed. F. rubra and T. pratense responded differently in compensatory growth and competition intensity to environmental change and co-existing species. The differences in their physiological and ecological traits may account for species-dependent responses. The present study emphasizes that predicting the plant assemblage response in the face of global change requires in understanding the integrating effects of abiotic and biotic factors.     

Can CO2 enrichment modify the effect of water and high light stress on biomass allocation and relative growth rate of cork oak seedlings?

To test whether the impact of an enriched-CO₂ environment on the growth and biomass allocation of first-season Quercus suber L. seedlings can modify the drought response under shade or sun conditions, seedlings were grown in pots at two CO₂ concentrations × two watering regimes × two irradiances. Compared to CO₂, light and water treatment had greater effects on all morphological traits measured (height, stem diameter, number of leaves, leaf area, biomass fractions). Cork oak showed particularly large increases in biomass in response to elevated CO₂ under low-watered (W−) and high-illuminated conditions (L+). Allocation shifted from shoot to root under increasing irradiance (L+), but was not affected by CO₂. Changes in allocation related to water limitation were only modest, and changed over time. Relative growth rate (RGR) and net assimilation rate (NAR) were significantly greatest in the L+/W+ treatment for both CO₂ concentrations. Changes in RGR were mainly due to NAR. Growth responses to increased light, water or CO₂ were strongest with light, medium with water availability and smallest for CO₂, in terms of RGR. The rise in NAR for light and water treatments was counterbalanced by a decrease in SLA (specific leaf area) and LMF (leaf mass fraction). Results suggest that elevated CO₂ caused cork oak seedlings to improve their performance in dry and high light environments to a greater extent than in well-irrigated and low light ones, thus ameliorating the effects of soil water stress and high light loads on growth.     

外部干扰对根茎型克隆植物甘草自然种群植株生长及主要药用成分含量的影响

克隆植物, 特别是游击型克隆植物, 其分株在一定时间段往往通过连接结构(即间隔子)连接在一起形成克隆网络, 基于克隆网络的克隆植物特性显著提高了克隆植物应对外部干扰的能力。长期封育禁牧使得甘草(Glycyrrhiza uralensis)自然种群免受外部干扰, 却并没有促使受破坏的甘草自然种群快速恢复。因为克隆整合、克隆储存及克隆分株的选择性放置等克隆植物特性的存在, 适度的外部干扰有利于根茎型克隆植物甘草的生长与繁衍。但外部干扰与克隆植物特性在甘草种群恢复中的作用及其机理目前并不清楚。该研究以甘草自然种群为研究对象, 通过野外控制实验进行模拟动物采食和人工采挖干扰, 探讨不同程度干扰对甘草自然种群生长和主要药用成分含量的影响。结果表明在中度放牧采食干扰或者人工采挖干扰下, 甘草自然种群的分株密度、高度和地上生物量与对照相比均没有显著差异, 甘草表现出等补偿生长; 而在不同类型的重度干扰下, 甘草产生的响应不一致。重度放牧采食干扰下甘草表现出欠补偿生长, 而重度人工采挖干扰下甘草表现出超补偿生长。克隆植物特性在甘草应对外部干扰过程中发挥着重要作用。外部干扰一定程度上提高了甘草中甘草苷和甘草酸的含量, 从而提升了药材的品质。进一步的研究应该关注于甘草自然种群与环境因子之间的关系, 解析甘草自然种群退化的原因, 提出相应的复壮技术, 探讨可持续利用的管理模式。     

Effect of defoliation intensity on aboveground and belowground relative growth rates

According to a simple growth model, grazed and ungrazed plants may have equal absolute growth rates provided that the relative growth rate (RGR) of grazed plants increases exponentially with grazing intensity (proportion of biomass removed). This paper reports results from an experiment designed to determine whether plants of two grass species subjected to a gradient of defoliation intensities, from 0 to 100% aboveground biomass removal, showed such a response. The relationship between aboveground RGR and defoliation intensity was exponential and closely matched the theoretical relationship of equal absolute growth rate. Thus, plants showed the same aboveground growth regardless of defoliation intensity thanks to an exponential stimulation of RGR by defoliation. Belowground RGR was depressed by defoliation of more than 20% of the above-ground biomass. In spite of the drastic modification imposed by the treatments on the relative proportions of different plant parts, after a 42-day recovery period basic allometric relationships, such as root:shoot and leafarea: weight ratios, were not affected by defoliation intensity. Exponential aboveground compensatory responses represent a key feedback process resulting in constant aboveground growth regardless of defoliation intensity and appear to be a simple consequence of strong commitments to certain allometric relationships.