环青海湖地区天然草地和退耕恢复草地植物群落生物量对氮、磷添加的响应

植物群落生物量反映了植被的初级生产能力, 是陆地生态系统碳(C)输入的最主要来源, 往往受到自然界中氮(N)、磷(P)元素供应的限制。该试验以青藏高原环青海湖地区的高寒草原为研究对象, 探讨了天然草地和退耕恢复草地植被群落生物量对N (10 g·m^-2)、P (5 g·m^-2)养分添加的响应。N、P添加显著增加了天然草地禾草的生物量, 进而促使地上总生物量显著提高。退耕恢复草地禾草和杂类草的生物量对N添加均有一致的正响应, 从而促使地上总生物量显著增加174%, 群落地上和地下总生物量显著增加34%; 而P添加对恢复草地生物量各项参数均无显著影响。回归分析显示: 天然草地植物群落地上生物量随土壤中NO₃^--N含量的增加而增加(p < 0.05), 退耕恢复草地植被地上、地下和总生物量均与土壤NO₃^--N含量显著正相关(p < 0.01), 说明环湖地区高寒草原植物生长主要受N供应的限制, P的限制作用随土地利用方式的转变和群落演替阶段的不同而变化; 相比天然草地, 恢复草地在现阶段植被初级生产力受N的限制作用更强烈, 土壤中可利用N含量是限制其植被自然恢复和重建的关键因子。     

The interactive effects of nitrogen addition and increased precipitation on gross ecosystem productivity in an alpine meadow

氮水添加对高寒草甸生态系统生产力的影响 降水变化和大气氮沉降增加对草原生态系统碳交换具有重要的影响,进而影响草地生产力、群落组成和生态系统功能。然而,氮水添加对高寒草甸生态系统碳交换的影响目前尚不清楚。因此,本研究在青藏高原高寒草甸布设氮水添加试验,设置4种不同处理：对照、 加氮、加水和同时添加氮水,对生态系统碳交换过程进行了连续4年的原位观测。研究结果发现,氮添加可以增加总生态系统生产力(GEP)、植物地上生物量、群落盖度和群落加权平均高度(CWMh),而水分添加没有显著影响。生态系统碳交换对氮水添加的响应在干湿年存在显著差异。水分添加仅在干旱年对净生态系统碳交换(NEE)具有显著影响,原因是GEP的增加量大于生态系统呼吸(ER)。相反,氮添加仅在湿润年显著提高了生态系统碳交换,其中GEP的增加归因于NEE的增加量大于ER。结构方程结果表明,氮添加主要通过增加优势种的盖度从而提高NEE。本研究强调了降水和优势物种在调节高寒草甸生态系统响应环境变化中的重要作用。     

青藏高原高寒草甸植被和土壤对短期禁牧的响应

为探讨青藏高原高寒草甸对短期禁牧的响应,设置冬季自由放牧和短期(2年)禁牧的对比试验。采用随机样方法调查植被群落盖度,分析地上和地下生物量、根冠比、植被地上和地下部分以及表层(0-10cm)土壤全碳、全氮和全磷含量、生态化学计量以及营养元素的关联性。研究结果显示:1)短期禁牧显著改变高寒草甸植被盖度、地上生物量、根冠比、植被全磷含量和N∶P,以及土壤全磷含量。2)相关性分析表明,禁牧后土壤全碳含量与植被地上全碳含量呈显著相关性,自由放牧后土壤全碳和全氮含量分别与植被地下部分全碳和全氮含量呈显著相关性。结果表明,不同的草原管理措施(禁牧、放牧)会改变高寒草甸植被与土壤养分分配及其平衡关系,同时,植被与表层土壤主要养分含量之间的关联性仅存在于部分植物器官与部分营养元素之间。     

青藏高原典型草地植被退化与土壤退化研究

采用野外样方调查和室内分析法,探讨了青藏高原不同退化程度高寒草原和高寒草甸植被群落结构、植物多样性、地上-地下生物量、根系分配及土壤理化特性差异。研究表明:(1)随着退化程度加剧,高寒草原禾草优势地位未改变,高寒草甸优势种莎草逐渐被杂类草取代。(2)随着退化程度加剧,高寒草原地上生物量显著降低(P0.05),高寒草甸地上生物量先保持稳定再下降。高寒草甸地下生物量较高寒草原地下生物量对退化响应更敏感。(3)高寒草原退化过程中,莎草地上物生量变化不明显(P0.05),禾草地上生物量贡献率由88.12%减少至53.54%,杂类草地上生物量贡献率由0.08%增加至42.81%;高寒草甸退化过程中,禾草和杂类草地上生物量先增加后减小,莎草地上生物量占比由69.15%减少至0.04%,杂类草地上生物量占比由12.56%增加至92.61%。(4)随着退化程度加剧,高寒草原根系向浅层迁移,高寒草甸根系向深层迁移。(5)退化对高寒草甸土壤含水量(θ)、土壤有机碳(SOC)、总氮(TN)及土壤容重(BD)影响均比高寒草原更强烈。本研究对青藏高原退化草地恢复治理具有重要的参考价值。     

增温施氮对高寒草甸生产力及生物量分配的影响

在青藏高原高寒草甸区设置模拟增温和氮添加处理,研究长期增温与外源氮输入对高寒草甸群落生产及其分配的影响.结果表明:开顶箱增温装置造成小环境暖干化,即显著提高地表空气温度1.6℃,提高表层土壤温度1.4℃,降低土壤含水量4.7%.2012、2013和2014年不施氮处理下增温分别降低地上生物量61.5%、108.8%和77.1%,在高氮(40和80kg N·hm^-2·a^-1)处理下增温对群落地上生物量无显著影响,这说明增温的影响依赖于氮添加水平,且施氮补偿了增温导致的土壤氮损失.增温导致根冠比增加,2012、2013和2014年不施氮处理下增温分别增加根冠比98.6%、60.7%和97.8%.在不增温处理下,植物群落地上、地下生物量的变化率均表现出低氮(10、20 kg N·hm^-2·a^-1)促进、高氮抑制的趋势,达到饱和阈值时的氮添加剂量分别为56.0和55.5 kg N·hm^-2·a^-1;而在增温处理下,地上、地下生物量随施氮量增加呈线性增加趋势.这说明增温改变了高寒草甸生物量分配对外源氮输入的响应模式,增温导致的土壤无机氮含量变化是生物量分配模式改变的主要原因.由氮添加试验估算的高寒草甸氮饱和阈值表明,高寒草甸对氮输入的敏感性高于其他类型草地.     

围封与放牧对青藏高原草地生物量与功能群结构的影响

以青藏高原高寒草甸、高寒草原、温性荒漠草原的退化草地为基础,比较了5年围封样地与放牧样地的生物量和群落结构。结果表明:(1)围封后3类草地地上总生物量较放牧样地分别显著增加了48.1%、10.8%、34.5%;地下生物量对围封的响应与地上总生物量一致,且围封后高寒草原0~10cm土层根系生物量比例较放牧地显著下降。(2)围封显著降低了高寒草甸的根冠比,高寒草原和温性荒漠草原无显著变化。(3)与放牧地相比,围封显著增加了高寒草甸和高寒草原禾本科植物的生物量比例,高寒草甸杂类草显著降低,温性荒漠草原功能群生物量比例无显著差异。     

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。     

氮素添加对青藏高原高寒草甸植物群落物种丰富度及其与地上生产力关系的影响

植物种群对有限资源的竞争是决定植物群落物种组成、多样性和生产力等群落结构和功能的主要因素。该文以青藏高原高寒草甸为研究对象, 研究了短期内不同水平的氮素添加对高寒草甸植物群落的影响。结果表明: 1)氮素添加提高了土壤中NO₃^--N等可利用资源的含量, 增加了植物群落植被的盖度, 减小了植被的透光率, 随着施氮量的增加, 群落中物种丰富度显著降低(p < 0.001); 2)氮素添加显著改变了植物群落的地上生产力(p < 0.05), 随着施氮量的增加, 地上生产力呈先增加后降低的变化趋势, 各功能群中禾草生物量显著增加, 而杂类草和豆科植物生物量随施氮量的增加逐渐减少; 3)物种多样性与植被透光率呈线性正相关(p < 0.05); 地上生产力与土壤NO₃^--N含量呈线性正相关(p < 0.05); 物种丰富度与地上生产力之间呈负相关关系。这说明短期内氮素添加通过改变土壤中NO₃^--N等可利用资源的含量而对植物群落物种组成和地上生产力产生影响。     

不同类型高寒草地群落结构与生产对施氮的响应及其敏感性

大气氮沉降增加被认为是目前重要的环境问题,会引起生物多样性的丧失和生态系统稳定性的降低。但作为草地改良的管理措施,养分添加被广泛应用于退化草地的恢复。但由于不同类型草地所处气候与群落组成的差异,对氮输入的响应可能不同。通过在藏北高原高寒草甸与高寒草甸草原设定长期氮添加梯度试验(对照, 25, 50, 100, 200 kg N hm~(-2) a~(-1)),来探讨氮输入对生物多样性与生产的影响,并估算不同类型高寒草地的氮饱和阈值。施氮对高寒草甸物种多样性指数无影响,而随着施氮量的提高高寒草甸草原植物物种数和多样性指数均逐渐降低。开始施肥前两年,随着施氮量提高高寒草甸地上生物量呈现逐渐增加趋势,随着施肥时间的延长地上生物量呈现先增加后降低的趋势。在高寒草甸草原随着施氮量提高地上生物量均呈现先增加后降低的趋势。随着施氮量提高,开始施氮前三年高寒草甸禾草植物地上生物量逐渐提高;随着施氮时间的延长,禾草和豆科植物地上生物量呈现先增加后降低的趋势。高寒草甸莎草植物地上生物量由施氮开始时的逐渐增加转变为先增加后降低趋势,最后变为逐渐降低的趋势,这说明施氮不利于莎草植物的生长。施氮只在施肥第四年显著提高杂草植物地上生物量。高寒草甸草原呈现不同的规律,开始施氮前三年随着施氮量提高,禾草植物地上生物量呈现先增加后降低的趋势;随着施氮时间的延长,禾草地上生物量逐渐提高。莎草和杂草植物地上生物量呈现先增加后降低趋势。利用对氮输入响应最敏感的植物功能群禾草生物量估算的高寒草甸和高寒草甸草原的氮饱和阈值分别是109.5、125.8 kg N hm~(-2) a~(-1),这说明高寒草甸氮敏感性显著高于高寒草甸草原。由此可见,未来氮沉降增加会对不同类型高寒草地产生不同的影响,在不同类型高寒草地进行施肥恢复时也应将氮饱和阈值的差异考虑在内。     

青藏高原高山嵩草高寒草甸不同退化阶段植物群落与土壤养分

选取西藏自治区拉萨市当雄县4块不同退化程度的高山嵩草高寒草甸,采用空间序列代替时间演替的方法,研究不同退化阶段高寒草甸的土壤理化性质和植物群落特征以及二者之间的相关关系。结果表明: 不同退化阶段高寒草甸的土壤有机碳、全氮、速效磷、速效钾、铵态氮、硝态氮和含水量均随土壤退化程度加剧呈降低的趋势,而pH值呈现升高的趋势。中度退化草甸的植物群落高度、丰富度指数、多样性指数、均匀度指数最大。群落盖度、总生物量均为未退化草甸最大、重度退化草甸最小。随着草甸退化程度加剧,莎草科生物量及比例下降,豆科和杂类草生物量及比例增加,禾本科生物量及比例先增加后减小;草甸植物群落地上生物量与土壤有机碳、全氮、全磷含量和土壤含水量呈显著正相关,与土壤pH值呈显著负相关。随着草甸植被的退化,土壤退化加重,最终表现为草地生产力显著下降。     

Hierarchical responses of plant stoichiometry to phosphorus addition in an alpine meadow Community北大核心CSCD

Aims Terrestrial carbon (C), nitrogen (N), phosphorus (P) stoichiometry will reflect the effects of adjustment to local growth conditions as well as species' replacements. However, it remains unclear about the hierarchical responses of plant C:N:P to P addition at levels of species and functional groups in the N-limited alpine meadow. Methods A field experiment of P enrichment was conducted in an alpine meadow on the Qinghai-Xizang Plateau during 2009-2013. The stoichiometric patterns of four functional groups (grass, sedge, legume and forb) and five representative species, Elymus nutans (grass), Kobresia humilis (sedge), Oxytropis ochrocephala (legume), Taraxacum lugubre (rosette forb), Geranium pylzowianum (upright forb) were investigated in 2013, and the effects of P addition on species dominance and plant biomass were also analyzed. Important findings Both plant nutrition content and C:N:P varied significantly after five years' P addition, and the responses were consistent at species-And functional group (exemplar species excluded)-levels in the alpine meadow. P addition had neutral effect on C concentrations of grasses, sedges and forbs at both species-And functional group (exemplar species excluded)-levels. P fertilization increased plant P concentrations and thus decreased C:P and N:P of the four functional groups (exemplar species excluded) and the corresponding species. N concentrations significantly decreased and C:N increased in grasses and sedges after P addition, and the species-level responses were consistent with the functional group (exemplar species excluded) level. P addition significantly increased N contents and decreased C:N in Oxytropis ochrocephala, but had neutral effect on N contents and C:N at the functional group (exemplar species excluded) level of the legumes. While N contents and C:N in forbs responded to P addition differently at species and functional group (exemplar species excluded) levels. In the N-limited alpine meadow, species dominance of grasses increased gradually after P addition due to the increased N and P use efficiencies, while the biomass proportion of forbs decreased because of the lowered nutrition use efficiency. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Responses of plant community biomass to nitrogen and phosphorus additions in natural and restored grasslands around Qinghai Lake Basin

Aims Plant biomass reflects the primary productivity of community vegetation, and is the main resource of carbon input in the terrestrial ecosystem. It is usually limited by nitrogen (N) and phosphorus (P) availability in the soil. Alpine grassland around Qinghai Lake Basin has experienced extensive land-use changes due to the cultivation of native grassland and vegetation recovery on cropped land. In this experiment, two grassland types were chosen, natural alpine grassland (NG) and its adjacent restored grassland (RG), to determine the responses of plant community biomass to N and P additions with different land-use. Methods NH₄NO₃ and Ca(H₂PO₄)₂·H₂O were added in a completely randomized block design, with medium levels of 10 g N·m^-2 and 5 g P·m^-2. Soil NO₃^--N and available P contents, and the plant community biomass were measured in the two grasslands. Two-way ANOVA was used to determine the effects of nutrient additions on all measured indicators, and regression analysis was used to analyze the correlations between plant biomass and soil NO₃^--N and available P contents.Important findings Results showed: (1) N and P additions both increased grass biomass in the NG, and significantly elevated the total aboveground biomass, with the promoting effect of N addition higher than that of P addition; N addition significantly increased both grass and forb biomass in the RG, and markedly promoted the total aboveground biomass, while P addition had no effects on the functional groups and total aboveground biomass (p > 0.05). (2) N and P additions both had no effects on the belowground and total biomass in the NG, whereas N addition significantly increased the total biomass by 34% in the RG, which suggested that the effect of N limitation on the vegetation primary productivity was stronger in the RG at present stage. (3) The aboveground biomass in the NG increased with soil NO₃^--N content (p < 0.05), and the above- and below-ground as well as the total biomass were all positively correlated with soil NO₃^--N content in the RG (p < 0.01). These results indicated that the plant growth in alpine grassland around Qinghai Lake Basin was prone to N limitation, and the effect of P limitation changed with land-use. Soil available N might be the key limiting factor for vegetation restoration and reconstruction in the RG. The “Grain for Green” project (the land-use policy) and atmospheric N deposition are benefiting both plant growth and C accumulation in the alpine grassland ecosystem around Qinghai Lake Basin.     

Precipitation and nitrogen addition enhance biomass allocation to aboveground in an alpine steppe

There are two important allocation hypotheses in plant biomass allocation: allometric and isometric. We tested these two hypotheses in an alpine steppe using plant biomass allocation under nitrogen (N) addition and precipitation (Precip) changes at a community level. An in situ field manipulation experiment was conducted to examine the two hypotheses and the responses of the biomass to N addition (10 g N m^?2 y^?1) and altered Precip (±50% precipitation) in an alpine steppe on the Qinghai–Tibetan Plateau from 2013 to 2016. We found that the plant community biomass differed in its response to N addition and reduced Precip such that N addition significantly increased aboveground biomass (AGB), while reduced Precip significantly decreased AGB from 2014 to 2016. Moreover, reduced Precip enhanced deep soil belowground biomass (BGB). In the natural alpine steppe, the allocation between AGB and BGB was consistent with the isometric hypotheses. In contrast, N addition or altered Precip enhanced biomass allocation to aboveground, thus leading to allometric growth. More importantly, reduced Precip enhanced biomass allocation into deep soil. Our study provides insight into the responses of alpine steppes to global climate change by linking AGB and BGB allocation.     

氮肥添加对青藏高原高寒草甸6个群落优势种生态化学计量学特征的影响

以青藏高原高寒草甸为研究对象, 通过人工氮肥添加试验, 研究6个群落优势种在不同施氮(N)水平下叶片碳(C)、N、磷(P)元素含量的变化以及生态化学计量学特征。结果表明: 自然条件下, 6个物种叶片N、P质量浓度存在显著的差异, 表现为: 黄花棘豆(Oxytropis ochrocephala)最高, 为24.5和2.51 g·kg^-1, 其叶片N含量低于而P含量高于我国其他草地的豆科植物; 其余5个物种叶片N、P质量浓度分别为11.5-18.1和1.49-1.72 g·kg^-1, 嵩草(Kobresia myosuroides)叶片N含量最低, 垂穗披碱草(Elymus nutans)叶片P含量最低, 与我国其他区域的研究结果相比, 其叶片N和P含量均低于我国其他草地非豆科植物。随氮素添加量的增大, 6种群落优势种叶片的C和P含量保持不变; 其他5种植物叶片N含量显著增加, 黄花棘豆叶片N含量保持不变。未添加氮肥时, 6种植物叶片N:P为7.3-11.2, 说明该区植物生长更多地受N限制。随N添加量的增加, 除黄花棘豆外, 其他5种植物叶片N:P大于16, 表现为植物生长受P限制。综合研究表明, 青藏草原高寒草甸植物叶片N含量较低, 植物受N影响显著, 但不同物种对N的添加反应不同, 豆科植物黄花棘豆叶片对N添加不敏感, 其他5个物种叶片全N含量随着N添加量的升高而增加, 该研究结果可为高寒草甸科学施肥提供理论依据。     

Response of Alpine Plants to Nitrogen Addition on the Tibetan Plateau: A Meta-analysis

To identify the general effects of nitrogen addition on alpine plants, we used a meta-analysis approach to synthesize 599 observations from 51 studies on the Tibetan Plateau. Nitrogen addition significantly increased plant height by 19.0 %, plant biomass by 29.7 %, graminoid aboveground biomass by 89.8 %, and sedge aboveground biomass by 75.6 % but significantly decreased legume aboveground biomass by 34.5 %, forb aboveground biomass by 23.8 %, and species richness by 11.2 %. The effect of nitrogen addition on aboveground plant biomass and plant height increased with increasing the nitrogen addition rate. The effect of nitrogen addition on plant height increased with increasing mean annual precipitation but decreased with increasing mean annual temperature. Our findings suggested that the effect of nitrogen addition on alpine plants varied with plant functional types and nitrogen addition rate. In addition, climatic warming and precipitation changes may regulate the response of alpine plants to nitrogen addition on the Tibetan Plateau.     

Effects of N addition on ecological stoichiometric characteristics in six dominant plant species of alpine meadow on the Qinghai-Xizang Plateau,China

以青藏高原高寒草甸为研究对象, 通过人工氮肥添加试验, 研究6个群落优势种在不同施氮(N)水平下叶片碳(C)、N、磷(P)元素含量的变化以及生态化学计量学特征。结果表明: 自然条件下, 6个物种叶片N、P质量浓度存在显著的差异, 表现为: 黄花棘豆(Oxytropis ochrocephala)最高, 为24.5和2.51 g·kg^-1, 其叶片N含量低于而P含量高于我国其他草地的豆科植物; 其余5个物种叶片N、P质量浓度分别为11.5-18.1和1.49-1.72 g·kg^-1, 嵩草(Kobresia myosuroides)叶片N含量最低, 垂穗披碱草(Elymus nutans)叶片P含量最低, 与我国其他区域的研究结果相比, 其叶片N和P含量均低于我国其他草地非豆科植物。随氮素添加量的增大, 6种群落优势种叶片的C和P含量保持不变; 其他5种植物叶片N含量显著增加, 黄花棘豆叶片N含量保持不变。未添加氮肥时, 6种植物叶片N:P为7.3-11.2, 说明该区植物生长更多地受N限制。随N添加量的增加, 除黄花棘豆外, 其他5种植物叶片N:P大于16, 表现为植物生长受P限制。综合研究表明, 青藏草原高寒草甸植物叶片N含量较低, 植物受N影响显著, 但不同物种对N的添加反应不同, 豆科植物黄花棘豆叶片对N添加不敏感, 其他5个物种叶片全N含量随着N添加量的升高而增加, 该研究结果可为高寒草甸科学施肥提供理论依据。