藏北高寒草甸群落结构与物种组成对增温与施氮的响应

气候变暖和氮沉降增加作为全球环境问题,将严重影响陆地生态系统的结构与功能.研究发现,近几十年来青藏高原增温显著,其中冬季升温最明显.而已有的研究更多关注全年增温,对冬季增温研究较少.本文基于高寒草甸地区增温和氮素添加影响研究的不足,在青藏高原高寒草甸区开展模拟增温和氮添加试验,研究长期增温与氮添加对高寒草甸群落结构与物种组成的影响.试验布设于2010年7月,地点在西藏当雄高寒草甸区,共有3种增温方式:对照、全年增温、冬季增温;每种增温处理下设置5个氮素添加梯度:0、10、20、40、80 kg N·hm^-2·a^-1,系统研究气候变暖与氮添加对高寒草甸生态系统群落结构与物种组成的影响.结果表明: 2012—2014年,增温与施氮处理均显著影响群落总盖度:全年增温处理降低了群落总盖度;在不施氮处理下,冬季增温降低了群落盖度,但在施氮处理下,随着氮剂量的提高群落盖度逐渐升高.增温与施氮对不同功能群植物的影响不同,增温处理降低了禾草与莎草植物盖度,而施氮提高了禾草植物盖度.相关分析表明,植被群落总盖度与生长旺盛期土壤含水量呈正相关关系,推测在降雨较少的季节增温导致的土壤含水量降低是群落盖度降低的主要原因.半干旱区高寒草甸土壤水分主要受降雨的调控,未来气候变化情景下,降雨时空格局的改变会显著影响植被群落盖度及组成,且大气氮沉降的增加对植被群落的影响也依赖于降雨条件的变化.     

Short-term responses of an alpine meadow community to removal of a dominant species along a fertilization gradient

Aims The relationship between biodiversity and ecosystem functioning has intrigued ecologists for several decades, but the effect of loss of a dominant species on community structure and functioning along a nutrient gradient remains poorly understood. The aim of this paper was to test the effect of a dominant species on community structure and function by conducting a species removal experiment along a fertilization gradient.Methods We removed the population of a dominant species (Elymus nutans) in a long-term fertilization field in an alpine meadow on the Tibetan Plateau, China. Univariate general linear models were used to evaluate the effects of fertilization and removal on above-ground vegetation characteristics, including photosynthetically active radiation in the understory, species richness, Shannon–Weiner diversity index, Simpson's dominance index, above-ground biomass (including different functional groups) and seedling richness and density.Important findings Results revealed that after two plant growing seasons, there was no significant effect of the removal of a dominant species on species richness and diversity of the remaining vegetation, but the biomass of forbs and seedling recruitment were significantly increased indicative of the potential for long-term effects. Moreover, removal had a large effect at high fertilization levels, but little effect when fertilization levels were low. Our studies indicated that community response to loss of a dominant species was mainly dependent on resource availability and the remaining functional group identities. We also found seedling recruitment was usually more sensitive to the influence of competition of dominant species than the established vegetation in the short term.     

Response of barley seedlings to UV-B radiation as affected by NaCl

The response of barley seedlings, subjected to 150 mmol/L NaCl for 4 days at different light regimes (4 d in the light, 4 d in darkness and a 12 h light/dark cycle) before UV-B radiation was investigated. NaCl treatment resulted in a decrease of total chlorophyll content and an increase in H2O2, free proline and lipid peroxidation, as quantified by measurement of malondialdehyde. Significantly more proline was accumulated in the light than in darkness. The combination of UV-B and NaCl treatment produced an additive effect on most of the parameters studied. UV-B radiation reduced the chlorophyll/carotenoids ratio and photochemical efficiency of PSII as estimated by chlorophyll fluorescence. NaCl pre-exposure decreased H2O2 generation and lipid peroxidation and alleviated the inhibitory effect of UV-B on PSII activity. Proline accumulated under salt stress conditions might be one of the reasons for the observed tolerance of barley seedlings to UV-B radiation.     

Susceptibility of pollen to UV-B radiation: an assay of 34 taxa

Much of the ultraviolet-B radiation (UV-B) research on plants has concentrated on vegetative plant parts, and only a small fraction has dealt with the reproductive system. The present study analyzed pollen grains of 34 taxa germinated and grown under two levels of UV-B radiation (187 and 460 mW/m2) or no UV-B (control group). Visible radiation at 260 mmol/m/s was present in all treatments. Taxa included those with binucleate and trinucleate pollen types. We detected differences among species. A significant reduction in pollen germination occurred in only five species. Pollen tubes of >50% of the species showed significant reduction in length. Trinucleate pollen types were more likely to exhibit tube length reduction than the binucleate types. Proportionately more monocotyledonous species were sensitive to UV-B treatment than dicotyledonous species, and proportionately more wild species were sensitive than cultivated species and pollen collected from plants growing in the field were somewhat more sensitive than pollen collected from plants grown in the greenhouse. Species in which pollination occurred earlier in the season were more likely to be susceptible to UV-B radiation than those for which anthesis took place later in the season, suggesting a possible adaptation to UV-B radiation.     

Linking individual response to biotic interactions with community structure: a trait-based framework

1.  Due to species-specificity of the outcomes of biotic interactions, it is difficult to generalize from observed biotic interactions at the individual plant level to the effect of those interactions at the community level. To evaluate the importance of biotic interactions in shaping plant communities, it is necessary to understand how the outcomes of the complex interactions observed at the individual level can influence community structure.
2.  Here, we propose a trait-based framework that identifies and organises mechanisms affecting community structure (here described with relative abundances of plant functional traits – i.e. the distribution of trait values at the community level). We applied our approach to a single leaf trait, specific leaf area (SLA), to link individual responses to plant interactions with community structure (SLA distribution observed at the community level) and to test whether biotic interactions can predict the functional composition of subalpine grasslands. We evaluated the generality of our model through a cross-validation with a set of eight subalpine grasslands independent from the four fields used to build the model.
3.  We found that competition and facilitation were able to explain the functional composition of subalpine grasslands, and the relevant fitness components (survival or growth) explaining this link changed depending on the limiting resources. When soil water availability was limiting, positive plant-plant interactions acting on survival were able to explain community structure. In contrast, when no water limitation was observed competition acting on individual growth was the main driver of community structure.
4.  Our framework enables evaluation of the consequences of biotic interactions observed at individual level on community structure, thereby indicating when and where different types of plant-plant interactions are important.     

滇西北退化高寒草甸植物群落结构对刈割的响应

为探究滇西北不同退化级别高寒草甸植物群落结构对外界干扰的响应敏感性,以香格里拉市的典型高寒草甸为研究对象,于2018-2020年在三个退化梯度上（严重退化,S1;中度退化,S2;轻度退化,S3）开展控制刈割实验,进而分析草甸植物物种丰富度、群落组成相似性、群落复杂度和关键种的变化规律。结果表明：（1）刈割后,S1的物种丰富度显著增加（P<0.05）,S2和S3的物种丰富度未发生显著变化（P>0.05）;（2）相较于S2和S3,S1梯度的植物群落组成变化最大;（3） S1、S2和S3的植物群落复杂度在刈割后均呈先下降后增加的趋势,但S1的植物群落复杂度变化幅度高于S2和S3;（4）刈割导致各退化草甸植物群落的关键种发生了变化,2018、2019和2020年S1梯度的关键种在豆科（Leguminsae）和蔷薇科（Rosaceae）之间变化,S2梯度的关键种在禾本科（Gramineae）和菊科（Compositae）之间变化,S3梯度的关键种在蔷薇科（Rosaceae）、菊科（Compositae）和禾本科（Gramineae）之间转变。研究表明,滇西北高寒草甸植物物种丰富度、群落组成和群落复杂度对外界干扰响应的敏感性可能随退化加剧而上升,但群落关键种的响应过程较复杂。     

Linking leaf veins to growth and mortality rates: an example from a subtropical tree community

A fundamental goal in ecology is to link variation in species function to performance, but functional trait–performance investigations have had mixed success. This indicates that less commonly measured functional traits may more clearly elucidate trait–performance relationships. Despite the potential importance of leaf vein traits, which are expected to be related to resource delivery rates and photosynthetic capacity, there are few studies, which examine associations between these traits and demographic performance in communities. Here, we examined the associations between species traits including leaf venation traits and demographic rates (Relative Growth Rate, RGR and mortality) as well as the spatial distributions of traits along soil environment for 54 co‐occurring species in a subtropical forest. Size‐related changes in demographic rates were estimated using a hierarchical Bayesian approach. Next, Kendall's rank correlations were quantified between traits and estimated demographic rates at a given size and between traits and species‐average soil environment. Species with denser venation, smaller areoles, less succulent, or thinner leaves showed higher RGR for a wide range of size classes. Species with leaves of denser veins, larger area, cheaper construction costs or thinner, or low‐density wood were associated with high mortality rates only in small size classes. Lastly, contrary to our expectations, acquisitive traits were not related to resource‐rich edaphic conditions. This study shows that leaf vein traits are weakly, but significantly related to tree demographic performance together with other species traits. Because leaf traits associated with an acquisitive strategy such as denser venation, less succulence, and thinner leaves showed higher growth rate, but similar leaf traits were not associated with mortality, different pathways may shape species growth and survival. This study suggests that we are still not measuring some of key traits related to resource‐use strategies, which dictate the demography and distributions of species.     

Using functional trait diversity to infer community assembly mechanisms: an exclosure experiment as an example

AimsThe Qinghai-Tibetan Plateau has a mean altitude exceeding 4000 m and covers about 2.5 million km2. More than 60% of this area is alpine grassland. Exclosures have been widely used in this region to study the sustainable use of grassland resources. We used patterns of functional trait diversity to infer the effects of exclosures on community assembly in alpine meadows.     

亚高寒草甸植物群落种多度分布关系及相似性对氮磷添加的响应

以青藏高原亚高寒草甸为研究对象,采用随机区组设计,通过连续4a添加N、P,研究了不同施肥(N、P、N+P)处理下群落物种丰富度、种多度分布模式以及群落相似性的变化特征。结果显示:(1)N、N+P连续添加4年后,随N素添加水平的增加,草地植物群落物种丰富度逐渐降低(P0.001);种多度分布曲线的斜率逐渐增大;N+P添加处理对植物群落物种丰富度和种多度分布(SAD)曲线的影响较单独N添加处理更显著,如N15P15处理下群落物种丰富度的降幅最大,达对照群落的65.5%;(2)单一N或N+P处理中,不同添加量间的植被组成趋异,而相同添加量的植被组成趋同(stress level=0.152);(3)N、N+P添加引起刷状根的丛生型禾本科植物逐渐在植物群落中占据优势;(4)P素添加对群落物种丰富度、种多度分布曲线、群落相似性和不同生长型组成及比例的影响不显著;(5)植物生长型特征和N/P添加处理可解释56.97%植物群落的物种多度分布特征。这些结果表明:亚高寒草甸地区N添加引起植物群落组成的重新排序、优势种的变化、SAD曲线逐渐陡峭,群落的相似性增加;N富集时,添加P素会增加N素的利用效率,且群落结构受N、P供应水平的影响。     

Plant community responses to increased precipitation and belowground litter addition: Evidence from a 5‐year semiarid grassland experiment

Global climate change is predicted to stimulate primary production and consequently increases litter inputs. Changing precipitation regimes together with enhanced litter inputs may affect plant community composition and structure, with consequent influence on diversity and ecosystem functioning. Responses of plant community to increased precipitation and belowground litter addition were examined lasting 5 years in a semiarid temperate grassland of northeastern China. Increased precipitation enhanced community species richness and abundance of annuals by 16.8% and 44%, but litter addition suppressed them by 25% and 54.5% after 5 years, respectively. During the study period, perennial rhizome grasses and forbs had consistent negative relationship under ambient plots, whereas positive relationship between the two functional groups was found under litter addition plots after 5 years. In addition, increased precipitation and litter addition showed significant interaction on community composition, because litter addition significantly increased biomass and abundance of rhizome grasses under increased precipitation plots but had no effect under ambient precipitation levels. Our findings emphasize the importance of water availability in modulating the responses of plants community to potentially enhanced litter inputs in the semiarid temperate grassland.     

A trait-based approach to community assembly: partitioning of species trait values into within- and among-community components

Plant functional traits vary both along environmental gradients and among species occupying similar conditions, creating a challenge for the synthesis of functional and community ecology. We present a trait-based approach that provides an additive decomposition of species' trait values into alpha and beta components: beta values refer to a species' position along a gradient defined by community-level mean trait values; alpha values are the difference between a species' trait values and the mean of co-occurring taxa. In woody plant communities of coastal California, beta trait values for specific leaf area, leaf size, wood density and maximum height all covary strongly, reflecting species distributions across a gradient of soil moisture availability. Alpha values, on the other hand, are generally not significantly correlated, suggesting several independent axes of differentiation within communities. This trait-based framework provides a novel approach to integrate functional ecology and gradient analysis with community ecology and coexistence theory.     

Plant community responses to nitrogen addition and increased precipitation: the importance of water availability and species traits

Global nitrogen (N) enrichment and changing precipitation regimes are likely to alter plant community structure and composition, with consequent influences on biodiversity and ecosystem functioning. Responses of plant community structure and composition to N addition and increased precipitation were examined in a temperate steppe in northern China. Increased precipitation and N addition stimulated and suppressed community species richness, respectively, across 6 years (2005–2010) of the manipulative experiment. N addition and increased precipitation significantly altered plant community structure and composition at functional groups levels. The significant relationship between species richness and soil moisture (SM) suggests that plant community structure is mediated by water under changing environmental conditions. In addition, plant height played an important role in affecting the responses of plant communities to N addition, and the effects of increased precipitation on plant community were dependent on species rooting depth. Our results highlight the importance and complexity of both abiotic (SM) and biotic factors (species traits) in structuring plant community under changing environmental scenarios. These findings indicate that knowledge of species traits can contribute to mechanistic understanding and projection of vegetation dynamics in response to future environmental change.     

Species richness loss after nutrient addition as affected by N:C ratios and phytohormone GA3 contents in an alpine meadow community

Aims From the light-competition hypothesis, competition for light is asymmetric and the observed increases in plant-size variability with increasingly denser canopies are primarily due to competition for light. Greater plant height provides pre-emptive access to light and produces increased height differences among species. The question is what produces these differences in plant height or height growth response among species in response to fertilization.Methods In 2009, a field experiment of N, P and N + P enrichments at three levels each was initiated in an alpine meadow on the northeast Qinghai-Tibet Plateau. Effects of fertilization on species richness, aboveground net primary production (ANPP), relative light intensity and plant height of different plant functional groups were determined. Festuca ovina (grass), Kobresia humilis (sedge), Oxytropis ochrocephala (legume), Taraxacum lugubre (rosette forb) and Geranium pylzowianum (upright forb) were selected as exemplars of each of the indicated functional groups. The N:C ratios in aboveground biomass, gibberellic acid (GA 3) concentrations in leaves, plant heights and height relative growth rate (RGR) of these exemplar species were analyzed in detail.Important findings Species richness of grasses significantly increased with increasing N + P levels. Species richness of legumes and upright forbs decreased after N and N + P additions. P addition had no significant effect on species richness. The effects of N + P addition on species richness and ANPP were consistently stronger than those of the single N or P fertilization. Reductions in species richness caused by nutrient addition paralleled the increases in ANPP and decreases in light intensity under the canopies, indicating indirect effect of nutrient addition on species richness via ANPP-induced light competition. The exemplar species that responded most positively to fertilization in height and RGR also displayed stronger increases in their GA 3 content and N:C ratios. GA 3 concentrations and N:C ratios were positively correlated with height RGR when the data were pooled for all species. The tallest and the fastest-growing grass, F. ovina, had the largest increase in N:C ratios and the highest leaf GA 3 concentrations after nutrient addition. These results indicated that differential responses of GA 3 concentrations and N:C ratios to fertilization were related to the inequality in plant heights among species.     

Ecological index of maturity to evaluate the vegetation disturbance of areas affected by restoration work: a practical example of its application in an area of the Southern Alps

This paper presents the ecological index of maturity (EIM), a new index for evaluating the vegetation disturbance of an area affected by environmental restoration measures based on the study of plant communities. The EIM is the result of the unification and development of some floristic‐vegetational indices devised by Taffetani and Rismondo (2009) and Rismondo et al. (2011) to assess the ecological functionality of agro‐ecosystems. The EIM provides a value between 0 (high vegetation disturbance) and 9 (undisturbed vegetation), obtained by considering three distinct variables of the species of a plant community: phytosociological class, chorotype, and coverage. Starting from the measure of the degree of maturity of the vegetation (dynamic stage of succession), EIM values tend to decrease the greater the coverage of exotic species and the lower the coverage of endemic species present in the plant community. A practical example, in which the EIM is applied to a mountainous area in the Southern Alps (Northern Italy) that, following a landslide, was subject to soil bioengineering work aimed at slope stabilization and environmental restoration, is provided in order to understand the importance and features of the EIM.     

Ambient UV-B radiation reduces PSII performance and net photosynthesis in high Arctic Salix arctica

Ambient ultraviolet-B (UV-B) radiation potentially impacts the photosynthetic performance of high Arctic plants. We conducted an UV-B exclusion experiment in a dwarf shrub heath in NE Greenland (74°N), with open control, filter control, UV-B filtering and UV-AB filtering, all in combination with leaf angle control. Two sites with natural leaf positions had ground angles of 0° (‘level site’) and 45° (‘sloping site’), while at a third site the leaves were fixed in an angle of 45° to homogenize the irradiance dose (‘fixed leaf angle site’). The photosynthetic performance of the leaves was characterized by simultaneous gas exchange and chlorophyll fluorescence measurements and the PSII performance through the growing season was investigated with fluorescence measurements. Leaf harvest towards the end of the growing season was done to determine the specific leaf area and the content of carbon, nitrogen and UV-B absorbing compounds. Compared to a 60% reduced UV-B irradiance, the ambient solar UV-B reduced net photosynthesis in Salix arctica leaves fixed in the 45° position which exposed leaves to maximum natural irradiance. Also a reduced Calvin Cycle capacity was found, i.e. the maximum rate of electron transport (J_max) and the maximum carboxylation rate of Rubisco (V_cmax), and the PSII performance showed a decreased quantum yield and increased energy dissipation. A parallel response pattern and reduced PSII performance at all three sites indicate that these responses take place in all leaves across position in the vegetation. These findings add to the evidence that the ambient solar UV-B currently is a significant stress factor for plants in high Arctic Greenland.