Plant community structure regulates responses of prairie soil respiration to decadal experimental warming

Soil respiration is recognized to be influenced by temperature, moisture, and ecosystem production. However, little is known about how plant community structure regulates responses of soil respiration to climate change. Here, we used a 13‐year field warming experiment to explore the mechanisms underlying plant community regulation on feedbacks of soil respiration to climate change in a tallgrass prairie in Oklahoma, USA. Infrared heaters were used to elevate temperature about 2 °C since November 1999. Annual clipping was used to mimic hay harvest. Our results showed that experimental warming significantly increased soil respiration approximately from 10% in the first 7 years (2000–2006) to 30% in the next 6 years (2007–2012). The two‐stage warming stimulation of soil respiration was closely related to warming‐induced increases in ecosystem production over the years. Moreover, we found that across the 13 years, warming‐induced increases in soil respiration were positively affected by the proportion of aboveground net primary production (ANPP) contributed by C₃ forbs. Functional composition of the plant community regulated warming‐induced increases in soil respiration through the quantity and quality of organic matter inputs to soil and the amount of photosynthetic carbon (C) allocated belowground. Clipping, the interaction of clipping with warming, and warming‐induced changes in soil temperature and moisture all had little effect on soil respiration over the years (all P > 0.05). Our results suggest that climate warming may drive an increase in soil respiration through altering composition of plant communities in grassland ecosystems.     

凋落物多样性及组成对凋落物分解和土壤微生物群落的影响

凋落物分解是生态系统营养物质循环的核心过程,而土壤微生物群落在凋落物分解过程中扮演着极其重要且不可替代的角色。随着生物多样性的丧失日益严峻,探讨凋落物多样性及组成对凋落物分解和土壤微生物群落的影响,不仅有助于了解凋落物分解的内在机制,而且可为退化草原生态系统的恢复提供参考。以内蒙古呼伦贝尔草原退化恢复群落中的草本植物为研究对象,依据植物多度、盖度、频度和物种的重要值及其在群落中的恢复程度筛选出排序前4的羊草（Leymus chinensis）、茵陈蒿（Artemisia capillaris）、麻花头（Serratula centauroides）、二裂委陵菜（Potentilla bifurca）的凋落物为实验材料,通过设置3种凋落物多样性水平（1,2,4）,包括11种凋落物组合（单物种凋落物共4种,两物种凋落物混合共6种,四物种凋落物混合共1种）,利用磷脂脂肪酸（PLFA）方法来研究分解60 d后凋落物多样性及组成对凋落物分解和土壤微生物群落的影响。结果表明：（1）凋落物物种多样性仅对C残余率具有显著影响,表现在两物种混合凋落物C残余率显著低于单物种凋落物,而凋落物组成对所观测的4个凋落物分解参数（质量、C、N残余率以及C/N）均具有显著影响;（2）凋落物物种多样性对细菌（B）含量具有显著影响,而凋落物组成对真菌（F）含量具有显著影响,两者对F/B以及微生物总量均无显著影响;（3）冗余分析结果表明凋落物组成与凋落物分解相关指标（凋落物质量、C、N残余率及C/N）和土壤微生物（真菌、细菌含量）的相关关系高于凋落物多样性。（4）进一步建立结构方程模型（Structural Equation Model,SEM）发现,凋落物初始C含量对凋落物质量、C、N残余率及C/N有显著正的直接影响;凋落物木质素含量对凋落物质量、C、N残余率有显著正的直接影响;凋落物初始N含量对N残余率有显著正的直接影响,而对C残余率及C/N有显著负的直接影响;凋落物初始C/N对凋落物质量、N残余率有显著正的直接影响,而对C/N有显著负的直接影响。此外,凋落物初始C、N、木质素含量及C/N均对真菌含量具有显著正影响,并可通过真菌对凋落物质量分解产生显著负的间接影响。以上结果表明该退化恢复区域优势种凋落物分解以初始C、木质素为主导,主要通过土壤真菌影响凋落物的分解进程,这将减缓凋落物的分解速率进而减慢草原生态系统的进程。这些结果为凋落物多样性及组成对自身分解和土壤微生物群落的影响提供了实验依据,也为进一步分析凋落物分解内在机制以及草原生态系统的恢复提供了数据参考。     

Plant diversity drives soil microbial biomass carbon in grasslands irrespective of global environmental change factors

Soil microbial biomass is a key determinant of carbon dynamics in the soil. Several studies have shown that soil microbial biomass significantly increases with plant species diversity, but it remains unclear whether plant species diversity can also stabilize soil microbial biomass in a changing environment. This question is particularly relevant as many global environmental change (GEC) factors, such as drought and nutrient enrichment, have been shown to reduce soil microbial biomass. Experiments with orthogonal manipulations of plant diversity and GEC factors can provide insights whether plant diversity can attenuate such detrimental effects on soil microbial biomass. Here, we present the analysis of 12 different studies with 14 unique orthogonal plant diversity × GEC manipulations in grasslands, where plant diversity and at least one GEC factor (elevated CO₂, nutrient enrichment, drought, earthworm presence, or warming) were manipulated. Our results show that higher plant diversity significantly enhances soil microbial biomass with the strongest effects in long‐term field experiments. In contrast, GEC factors had inconsistent effects with only drought having a significant negative effect. Importantly, we report consistent non‐significant effects for all 14 interactions between plant diversity and GEC factors, which indicates a limited potential of plant diversity to attenuate the effects of GEC factors on soil microbial biomass. We highlight that plant diversity is a major determinant of soil microbial biomass in experimental grasslands that can influence soil carbon dynamics irrespective of GEC.     

植物、土壤及土壤管理对土壤微生物群落结构的影响

土壤微生物是土壤生态系统的重要组成部分,对土壤微生物群落结构多样性的研究是近年来土壤生态学研究的热点。本文综述了有关植物、土壤类型以及土壤管理措施对土壤微生物群落结构影响的最新研究结果,指出植物的作用因植物群落结构多样性、植物种类、同种植物不同的基因型,甚至同一植物不同根的区域而异;而土壤的作用与土壤质地和有机质含量等因素有关;植物和土壤类型在对土壤微生物群落结构影响上的作用存在互作关系。不同的土壤管理措施对土壤微生物群落结构影响较大,长期连作、大量的外援化学物质的应用降低了土壤微生物的多样性;而施用有机肥、免耕可以增加土壤微生物群落结构多样性,有利于维持土壤生态系统的功能。     

Plant diversity predicts beta but not alpha diversity of soil microbes across grasslands worldwide

Aboveground–belowground interactions exert critical controls on the composition and function of terrestrial ecosystems, yet the fundamental relationships between plant diversity and soil microbial diversity remain elusive. Theory predicts predominantly positive associations but tests within single sites have shown variable relationships, and associations between plant and microbial diversity across broad spatial scales remain largely unexplored. We compared the diversity of plant, bacterial, archaeal and fungal communities in one hundred and forty‐five 1 m² plots across 25 temperate grassland sites from four continents. Across sites, the plant alpha diversity patterns were poorly related to those observed for any soil microbial group. However, plant beta diversity (compositional dissimilarity between sites) was significantly correlated with the beta diversity of bacterial and fungal communities, even after controlling for environmental factors. Thus, across a global range of temperate grasslands, plant diversity can predict patterns in the composition of soil microbial communities, but not patterns in alpha diversity.     

Broad-scale analysis of soil microbial community DNA from Upland grasslands

We have applied a broad-scale approach to the analysis of DNA extracted from soils which support characteristic grasslands at an upland site in the UK. To test for the degree of coherence between microbial and vascular communities, grasslands were characterised as improved, semi-improved, or unimproved, according to the degree of management they had received and consequent botanical composition. Microbial DNA was extracted directly from the grassland soils and analysed by three techniques: (i) thermal denaturation, which profiles the guanine and cytosine (G÷C) base distribution within the community; (ii) cross hybridisation of the DNA which measures the degree of similarity between the samples; (iii) measurement of reassociation kinetics of denatured DNA, which provides a measure of the complexity of the DNA. Thermal denaturation revealed significant differences in the %G+C composition of the communities. DNA from the improved soil had the highest median %G+C value, whilst that from the unimproved soil had the lowest. The relative distribution of G+C bases also differed significantly between the samples from the three grasslands. Cross hybridisation of DNA from the different soils also indicated significant differences in the degree of similarity between the DNA from the grasslands, with unimproved showing 59% similarity to improved. Indices from the cross hybridisation assay suggested that, in terms of complexity, the samples ranked unimproved > semi-improved > improved. Reassociation kinetics supported this conclusion, but the rates of reassociation were such that less than 40% reassociation occurred over a 31-day period, thus preventing calculation of C _o t_1/2.     

Plant presence and species combination, but not diversity, influence denitrifier activity and the composition of nirK-type denitrifier communities in grassland soil

To explore potential links between plant communities, soil denitrifiers and denitrifier function, the impact of presence, diversity (i.e. species richness) and plant combination on nirK -type denitrifier community composition and on denitrifier activity was studied in artificial grassland plant assemblages over two consecutive years. Mesocosms containing zero, four and eight species and different combinations of two species were set up. Differences in denitrifier community composition were analysed by canonical correspondence analyses following terminal restriction fragment length polymorphism analysis of PCR-amplified nirK gene fragments coding for the copper-containing nitrite reductase. As a measure of denitrifier function, denitrifier enzyme activity (DEA) was determined in the soil samples. The presence as well as the combination of plants and sampling time, but not plant diversity, affected the composition of the nirK -type denitrifier community and DEA. Denitrifier activity significantly increased in the presence of plants, especially when they were growing during summer and autumn. Overall, we found a strong and direct linkage of denitrifier community composition and functioning, but also that plants had additional effects on denitrifier function that could not be solely explained by their effects on nirK -type denitrifier community composition.     

Plant functional groups mediate effects of climate and soil factors on species richness and community biomass in grasslands of Mongolian Plateau

植物功能群在调控气候和土壤因子对蒙古高原草原群落物种丰富度和生物量影响中的作用 植物功能群组成主要受环境因素驱动,同时植物功能群组成也是影响草地生物多样性和生产力的主要因素之一。因此,理解植物功能群在调控环境因素对生态系统功能和生物多样性影响中可能发挥的作用至关重要。通过对蒙古高原草原65个样点的植物生物量和物种丰富度的调查,将157种多年生草本植物分为两种植物功能群(即禾草和杂类草)。通过随机森林模型和普通最小二乘回归,确定与植物功能群物种丰富度和地上生物量显著相关的环境因素(即干燥度、土壤总氮和pH),并利用结构方程模型探讨筛选出的环境因素与群落物种丰富度和生物量间的关系,以及植物功能群在驱动这种关系中发挥的作用。干燥度与禾草、杂类草以及整个群落的地上生物量和物种丰富度均呈显著的单峰关系。所有的物种丰富度和生物量指标均与土壤总氮和pH值显著相关。禾草在维持蒙古高原草原生态系统群落生物量中起着关键作用,并受气候因素的直接影响。而杂类草物种丰富度决定了群落总丰富度,并受到土壤因素直接的调控。因此,群落组成在调控环境因素对群落生物量和植物多样性的影响中起着关键作用。     

草地土壤呼吸研究进展

土壤呼吸是生态系统物质循环和能量流动的重要过程,具有多方面的生态意义。在综合比较了土壤呼吸的各种测定方法,概括了草地土壤呼吸的速率和年排放量,并就影响草地土壤呼吸的一些重要因素进行了阐述,文章最后讨论了草地土壤呼吸今后的研究方向。     

Relationship between plant species diversity and soil microbial functional diversity along a longitudinal gradient in temperate grasslands of Hulunbeir,Inner Mongolia,China

Numerous experiments have been established to examine the effect of plant diversity on the soil microbial community. However, the relationship between plant diversity and microbial functional diversity along broad spatial gradients at a large scale is still unexplored. In this paper, we examined the relationship of plant species diversity with soil microbial biomass C, microbial catabolic activity, catabolic diversity and catabolic richness along a longitudinal gradient in temperate grasslands of Hulunbeir, Inner Mongolia, China. Preliminary detrended correspondence analysis (DCA) indicated that plant composition showed a significant separation along the axis 1, and axis 1 explained the main portion of variability in the data set. Moreover, DCA-axis 1 was significantly correlated with soil microbial biomass C (r = 0.735, P = 0.001), microbial catabolic activity (average well color development; r = 0.775, P < 0.001) and microbial functional diversity (catabolic diversity: r = 0.791, P < 0.001 and catabolic richness: r = 0.812, P < 0.001), which suggested thatsome relationship existed between plant composition and the soil microbial community along the spatial gradient at a large scale. Soil microbial biomass C, microbial catabolic activity, catabolic diversity and catabolic richness showed a significant, linear increase with greater plant species richness. However, many responses that we observed could be explained by greater aboveground plant biomass associated with higher levels of plant diversity, which suggested that plant diversity impacted the soil microbial community mainly through increases in plant production.     

凋落物和根系去除对滨海沙地土壤微生物群落组成和功能的影响

通过在亚热带滨海防护林湿地松、尾巨桉、纹荚相思和木麻黄人工林中设置去除凋落物、去除根系和对照处理,分析改变地上、地下碳输入对沙地人工林土壤微生物生物量、群落结构和功能的影响.2015年9月,在处理设置一年后采集各处理样方0～10 cm土壤样品,分别采用磷脂脂肪酸分析方法和微孔板酶检测技术对土壤样品的微生物群落组成和6种酶活性进行测定.结果表明: 碳输入处理对土壤微生物生物量的影响因树种而异,并主要取决于凋落物和根系的质量.在尾巨桉林中,去除根系使得脂肪酸总量、革兰氏阳性细菌、革兰氏阴性细菌、真菌和放线菌生物量分别显著减少了31%、30%、32%、36%和26%,去除凋落物使得革兰氏阳性细菌、真菌和放线菌生物量显著减少了24%、27%和24%,而其他树种处理对微生物生物量无显著影响.碳输入处理对土壤微生物群落结构的影响主要表现为真菌丰度下降而放线菌丰度上升.不同树种的土壤酶活性对处理的响应表现不一致:湿地松、纹荚相思和木麻黄林分去除凋落物显著降低土壤中纤维素水解酶、β-葡萄糖苷酶、酸性磷酸酶和乙酰氨基葡萄糖苷酶活性,去除根系仅分别降低和提高了湿地松和纹荚相思林β-葡萄糖苷酶的活性;湿地松、木麻黄人工林去除凋落物显著降低了多酚氧化酶和过氧化物酶活性;去除根系对所有树种土壤氧化酶活性影响不显著.不同树种的凋落物、根系特性是影响土壤微生物群落组成和酶活性的重要因子,碳输入处理造成的土壤温度、水分等微环境的改变也是土壤微生物性质变化的重要驱动力.     

Short‐term harvesting of biomass from conservation grasslands maintains plant diversity

High yields are a priority in managing biomass for renewable energy, but the environmental impacts of various feedstocks and production systems should be equally considered. Mixed‐species, perennial grasslands enrolled in conservation programs are being considered as a source of biomass for renewable energy. Conservation grasslands are crucial in sustaining native biodiversity throughout the US Upper Midwest, and the effects of biomass harvest on biodiversity are largely unknown. We measured the effect of late‐season biomass harvest on plant community composition in conservation grasslands in three regions of Minnesota, USA from 2009 to 2012. Temporal trends in plant species composition within harvested grasslands were compared to unharvested grasslands using mixed effects models. A before‐after control‐impact approach using effect sizes was applied to focus on pre‐ and postharvest conditions. Production‐scale biomass harvest did not affect plant species richness, species or functional group diversity, nor change the relative abundance of the main plant functional groups. Differences in the relative abundances of plant functional groups were observed across locations; and at some locations, changed through time. The proportion of non‐native species remained constant, while the proportion of noxious weeds decreased through time in both harvested and unharvested grasslands at the central location. Ordination revealed patterns in species composition due to location, but not due to harvest treatment. Therefore, habitat and bioenergy characteristics related to grassland plant communities are not expected to change due to short‐term or intermittent late‐season biomass harvest.     

Plant communities and conservation of Botrychium-rich grasslands in the Bitcherland (Northern Vosges Biosphere Reserve, France)

Two species of Botrychium are present in the Bitcherland grasslands, B. lunaria and B. matricariifolium, and both are protected in this territory. They occur in five different continental grassland communities, (i) a dune grassland (Diantho-Armerietum), (ii) a mesotrophic sandy grassland (Viscario-Avenetum), composed of three sub-units, therophitic ( V.A. airetosum), meso-eutrophic (V.A. saxifragetosum) and acidic (V.A. violetosum caninae), and (iii) an oligotrophic sandy grassland (Aveno-Genistetum). Both species of Botrychium appear in the three sub-units of Viscario-Avenetum, but only one in Diantho-Armerietum (B. matricariifolium) and in Aveno-Genistetum (B. lunaria). These Botrychium species are present in quite similar habitats in other lowland areas in Central Europe. The conservation of these threatened species, sensitive to plant competition, requires the maintenance or restoration of extensive agricultural management with late cutting and no fertilization.     

新疆绿洲农田不同连作年限棉花根际土壤微生物群落多样性

以南北疆不同连作年限棉花根际土壤为研究对象,采用Biolog技术,并结合传统平板培养法和土壤酶的测定,研究连作对棉花根际土壤微生物群落多样性的影响。Biolog分析结果表明,不同连作年限棉花根际土壤微生物碳源利用和功能多样性差异显著。荒地土壤微生物活性较低;在连作年限较短时(5—10a),根际土壤微生物群落的平均颜色变化率(AWCD)和Shannon指数较高;长期连作(15—20a),则呈明显下降趋势。主成分分析表明,不同连作年限的棉花根际土壤微生物碳源利用特征有明显不同。第一、二组开垦与未开垦土壤分别在PC1和PC2上出现差异,未开垦土壤得分均为负值,开垦土壤均为正值;而正茬与连作多年的棉花土壤在PC1上差异显著。其中在PC1上起分异作用的碳源主要是羧酸类和聚合物类,这两类碳源可能是影响连作棉花根际土壤微生物的主要碳源。可培养微生物数量的测定结果表明,荒地细菌数量最少;在连作年限较低时(5—10a左右),细菌数量呈上升趋势;而长期连作(>15a)后,细菌数量呈现下降趋势。真菌数量在连作多年后(10—15a)也开始增加。放线菌变化趋势不明显。四种土壤酶活性在连作的初中期(5—15a),连作障碍表现明显,土壤酶活性呈下降(过氧化氢酶和磷酸酶)或先升高后下降(脲酶和蔗糖酶)趋势,但随着连作年限的延长(15—20a),这4种土壤酶活性均表现出增高趋势。综上所述,棉花长期连作使棉花根际土壤微生物群落多样性降低,发生连作障碍,进而导致棉花产量降低。     

Plant diversity loss reduces soil respiration across terrestrial ecosystems

The rapid global biodiversity loss has led to the decline in ecosystem function. Despite the critical importance of soil respiration (Rs) in the global carbon and nutrient cycles, how plant diversity loss affects Rs remains uncertain. Here we present a meta‐analysis using 446 paired observations from 95 published studies to evaluate the effects of plant and litter mixtures on Rs and its components. We found that total Rs and heterotrophic respiration (Rh) were, on average, greater in plant mixtures than expected from those of monocultures. These mixture effects increased with increasing species richness (SR) in both plant and litter mixtures. While the positive effects of species mixtures remained similar over time for total Rs, they increased over time for Rh in plant mixtures but decreased in litter mixtures. Despite the wide range of variations in mean annual temperature, annual aridity index, and ecosystem types, the plant mixture effects on total Rs and Rh did not change geographically, except for a more pronounced increase of total Rs in species mixtures with reduced water availability. Our structural equation models suggested that the positive effects of SR and stand age on total and Rh were driven by increased plant inputs and soil microbial biomass. Our results suggest that plant diversity loss has ubiquitous negative impacts on Rs, one of the fundamental carbon‐cycle processes sustaining terrestrial element cycling and ecosystem function.     

根际效应对大豆田土壤线虫群落组成及多样性的影响

根际作为重要的环境界面是植物与环境之间物质能量交换的场所,关于根际效应的研究已成为土壤生态学的新兴热点领域,然而有关大豆根际效应对土壤动物多样性影响的研究报道并不多见。在三江平原选择连续耕作15a的大豆田,对大豆根际区与非根际区土壤线虫群落结构组成进行了对比分析。结果表明:大豆根际区土壤线虫总数、辛普森多样性指数(Dom)显著高于非根际区,根际区的物种数(S)、物种丰富度指数(SR)显著低于非根际区。说明大豆根际效应增加土壤线虫的丰度,但降低了线虫群落结构的复杂性。大豆根际区植物寄生线虫(PP)、食真菌线虫(FF)和食细菌线虫(BF)数量显著高于非根际区,而PP类群的比例在根际区却显著低于非根际区。这一研究结果表明食微线虫(FF和BF)类群在大豆根际区的比例增加更显著。食真菌与食细菌线虫数量比值(F/B)指示大豆根际区细菌生物量相对高于真菌生物量。研究结果丰富了农田土壤线虫多样性的研究内容,并为我国东北大豆田线虫病害的防治及定制相应的农业管理措施提供参考。     

Plant functional group diversity promotes soil protist diversity

We tested whether effects of plant diversity can propagate through food webs, down to heterotrophic protists not linked directly to plants. To this end we synthesised grassland ecosystems with varying numbers of plant functional groups (FGN) and assessed corresponding changes in testate amoebae communities. The number of plant species was kept constant. When FGN was increased from 1 to 3, species number and total community density of live testate amoebae were enhanced according to a linear and a saturating function, respectively. From FGN 1 to 2, the appearance of new testate amoebae species did not affect the presence of the resident species, whereas, from FGN 2 to 3 about one quarter of the resident testate amoebae species was replaced, without altering the total species number. Overall, density by species increased, while evenness of the testate amoebae community was not affected by FGN; although Trinema lineare, one of the most common species, became more abundant. The observed relationship between plant functional group diversity and testate amoebae diversity could shed new light on the biogeographical distribution patterns of protists.     

Increases in soil respiration following labile carbon additions linked to rapid shifts in soil microbial community composition

Organic matter decomposition and soil CO₂ efflux are both mediated by soil microorganisms, but the potential effects of temporal variations in microbial community composition are not considered in most analytical models of these two important processes. However, inconsistent relationships between rates of heterotrophic soil respiration and abiotic factors, including temperature and moisture, suggest that microbial community composition may be an important regulator of soil organic matter (SOM) decomposition and CO₂ efflux. We performed a short-term (12-h) laboratory incubation experiment using tropical rain forest soil amended with either water (as a control) or dissolved organic matter (DOM) leached from native plant litter, and analyzed the effects of the treatments on soil respiration and microbial community composition. The latter was determined by constructing clone libraries of small-subunit ribosomal RNA genes (SSU rRNA) extracted from the soil at the end of the incubation experiment. In contrast to the subtle effects of adding water alone, additions of DOM caused a rapid and large increase in soil CO₂ flux. DOM-stimulated CO₂ fluxes also coincided with profound shifts in the abundance of certain members of the soil microbial community. Our results suggest that natural DOM inputs may drive high rates of soil respiration by stimulating an opportunistic subset of the soil bacterial community, particularly members of the Gammaproteobacteria and Firmicutes groups. Our experiment indicates that variations in microbial community composition may influence SOM decomposition and soil respiration rates, and emphasizes the need for in situ studies of how natural variations in microbial community composition regulate soil biogeochemical processes.     

Relationships between vegetation and bird community composition in grasslands of the Serengeti

The Serengeti–Mara ecosystem holds one of the largest natural grasslands of the world, which is well known for its large herds of mammals. However, the bird community structure of these grasslands has hardly ever been studied. For the first time, a large‐scale study on grassland bird communities has been conducted in all typical open grasslands of the Serengeti National Park. We used ten grassland plots representing a gradient of increasing vegetation height and also including shrubs and trees to analyse the influence of vegetation structure on the composition of grassland bird communities. Three communities of breeding birds were identified in relation to (a) short grass, (b) intermediate and long grass and (c) wooded grasslands. The bird communities of intermediate, long and wooded grassland were very similar, because of identical dominant bird species. Our results suggest that breeding birds of East African grasslands exhibit two contrasting habitat relationships: (1) birds that are restricted to short grass, and (2) species that are tolerant to vegetational shifts from intermediate grass via long grass to early stages of woody vegetation. Because the vegetation is often driven beyond this range in managed tropical grasslands through high grazing pressure or through shrub encroachment, we expect that the long‐grass bird communities will not generally resist anthropogenic disturbance.