Effects of six years atmospheric CO2 enrichment on plant,soil, and soil microbial C of a calcareous grassland

Stimulated plant production and often even larger stimulation of photosynthesis at elevated CO₂ raise the possibility of increased C storage in plants and soils. We analysed ecosystem C partitioning and soil C fluxes in calcareous grassland exposed to elevated CO₂ for 6 years. At elevated CO₂, C pools increased in plants (+23%) and surface litter (+24%), but were not altered in microbes and soil organic matter. Soils were fractionated into particle size and density separates. The amount of low-density macroorganic C, an indicator of particulate soil C inputs from root litter, was not affected by elevated CO₂. Incorporation of C fixed during the experiment (C_new) was tracked by C isotopic analysis of soil fractions which were labelled due to ¹³C depletion of the commercial CO₂ used for atmospheric enrichment. This data constrains estimates of C sequestration (absolute upper bound) and indicates where in soils potentially sequestered C is stored. C_new entered soils at an initial rate of 210±42 g C m^–2 year^–1, but only 554±39 g C_new m^–2 were recovered after 6 years due to the low mean residence time of 1.8 years. Previous process-oriented measurements did not indicate increased plant–soil C fluxes at elevated CO₂ in the same system (¹³C kinetics in soil microbes and fine roots after pulse labelling, and minirhizotron observations). Overall experimental evidence suggests that C storage under elevated CO₂ occurred only in rapidly turned-over fractions such as plants and detritus, and that potential extra soil C inputs were rapidly re-mineralised. We argue that this inference does not conflict with the observed increases in photosynthetic fixation at elevated CO₂, because these are not good predictors of plant growth and soil C fluxes for allometric reasons. C sequestration in this natural system may also be lower than suggested by plant biomass responses to elevated CO₂ because C storage may be limited by stabilisation of C_new in slowly turned-over soil fractions (a prerequisite for long-term storage) rather than by the magnitude of C inputs per se.     

Responses of soil microbiota of a late successional alpine grassland to long term CO2 enrichment

We investigated microbial responses in a late successional sedge-dominated alpine grassland to four seasons of CO₂ enrichment. Part of the plots received fertilizer equivalent to 4.5g N m⁻² a⁻¹. Soil basal respiration (R _mic ), the metabolic quotient for CO₂ (qCO₂=R _mic /C _mic ), microbial C and N (C _mic and N _mic ) as well as total soil organic C and N showed no response to CO₂ enrichment alone. However, when the CO₂ treatment was combined with fertilizer addition R _mic and qCO₂ were statistically significantly higher under elevated CO₂ than under ambient conditions (+57% and +71%, respectively). Fertilizer addition increased microbial N pools by 17%, but this was not influenced by elevated CO₂. Microbial C was neither affected by elevated CO₂ nor fertilizer. The lack of a CO₂-effect in unfertilized plots was suprising in the light of our evidence (based on C balance) that enhanced soil C inputs must have occurred under elevated CO₂ regardless of fertilizer treatment. Based on these data and other published work we suggest that microbial responses to elevated CO₂ in such stable, late-successional ecosystems are limited by the availability of mineral nutrients and that results obtained with fertile or heavily disturbed substrates are unsuitable to predict future microbial responses to elevated CO₂ in natural systems. However, when nutrient limitation is removed (e.g. by wet nitrogen deposition) microbes make use of the additional carbon introduced into the soil system. We believe that the response of natural ecosystems to elevated CO₂ must be studied in situ in natural, undisturbed systems.     

A field study of the effects of elevated CO2 on plant biomass and community structure in a calcareous grassland

The effects of elevated CO₂ on plant biomass and community structure have been studied for four seasons in a calcareous grassland in northwest Switzerland. This highly diverse, semi-natural plant community is dominated by the perennial grass Bromus erectus and is mown twice a year to maintain species composition. Plots of 1.3 m² were exposed to ambient or elevated CO₂ concentrations (n = 8) using a novel CO₂ exposure technique, screen-aided CO₂ control (SACC) starting in March 1994. In the 1st year of treatment, the annual harvested biomass (sum of aboveground biomass from mowings in June and October) was not significantly affected by elevated CO₂. However, biomass increased significantly at elevated CO₂ in the 2nd (+20%, P = 0.05), 3rd (+21%, P = 0.02) and 4th years (+29%, P = 0.02). There were no detectable differences in root biomass in the top 8 cm of soil between CO₂ treatments on eight out of nine sampling dates. There were significant differences in CO₂ responsiveness between functional groups (legumes, non-leguminous forbs, graminoids) in the 2nd (P = 0.07) and 3rd (P < 0.001) years of the study. The order of CO₂ responsiveness among functional groups changed substantially from the 2nd to the 3rd year; for example, non-leguminous forbs had the smallest relative response in the 2nd year and the largest in the 3rd year. By the 3rd year of CO₂ exposure, large species-specific differences in CO₂ response had developed. For five important species or genera the order of responsiveness was Lotus corniculatus (+271%), Carex flacca (+249%), Bromus erectus (+33%), Sanguisorba minor (no significant CO₂ effect), and six Trifolium species (a negative response that was not significant). The positive CO₂ responses in Bromus and Carex were most closely related to increases in tiller number. Species richness was not affected by CO₂ treatment, but species evenness increased under elevated CO₂ (modified Hill ratio; P = 0.03) in June of the 3rd year, resulting in a marginally significant increase in species diversity (Simpson's index; P = 0.09). This and other experiments with calcareous grassland plants show that elevated atmospheric CO₂ concentrations can substantially alter the structure of calcareous grassland communities and may increase plant community biomass. Received: 12 July 1997 / Accepted: 14 September 1998     

Effects of elevated CO2 and phosphorus addition on productivity and community composition of intact monoliths from calcareous grassland

We investigated the effects of elevated CO₂ (600 μl l⁻¹ vs 350 μl l⁻¹) and phosphorus supply (1 g P m⁻² year⁻¹ vs unfertilized) on intact monoliths from species-rich calcareous grassland in a greenhouse. Aboveground community dry mass remained almost unaffected by elevated CO₂ in the first year (+6%, n.s.), but was significantly stimulated by CO₂ enrichment in year two (+26%, P<0.01). Among functional groups, only graminoids contributed significantly to this increase. The effect of phosphorus alone on community biomass was small in both years and marginally significant only when analyzed with MANOVA (+6% in year one, +9% in year two, 0.1 ≥P > 0.05). Belowground biomass and stubble after two seasons were not different in elevated CO₂ and when P was added. The small initial increase in aboveground community biomass under elevated CO₂ is explained by the fact that some species, in particular Carex flacca, responded very positively right from the beginning, while others, especially the dominant Bromus erectus, responded negatively to CO₂ enrichment. Shifts in community composition towards more responsive species explain the much larger CO₂ response in the second year. These shifts, i.e., a decline in xerophytic elements (B. erectus) and an increase in mesophytic grasses and legumes occurred independently of treatments in all monoliths but were accelerated significantly by elevated CO₂. The difference in average biomass production at elevated compared to ambient CO₂ was higher when P was supplied (at the community level the CO₂ response was enhanced from 20% to 33% when P was added, in graminoids from 17% to 27%, in legumes from 4% to 60%, and in C. flacca from 120% to 298% by year two). Based on observations in this and similar studies, we suggest that interactions between CO₂ concentration, species presence, and nutrient availability will govern community responses to elevated CO₂. Received: 12 July 1997 / Accepted: 28 March 1998     

Nutrient relations in calcareous grassland under elevated CO2

Plant nutrient responses to 4 years of CO₂ enrichment were investigated in situ in calcareous grassland. Beginning in year 2, plant aboveground C:N ratios were increased by 9% to 22% at elevated CO₂ (P < 0.01), depending on year. Total amounts of N removed in biomass harvests during the first 4 years were not affected by elevated CO₂ (19.9 ± 1.3 and 21.1 ± 1.3 g N m⁻² at ambient and elevated CO₂), indicating that the observed plant biomass increases were solely attained by dilution of nutrients. Total aboveground P and tissue N:P ratios also were not altered by CO₂ enrichment (12.5 ± 2 g N g⁻¹ P in both treatments). In contrast to non-legumes (>98% of community aboveground biomass), legume C/N was not reduced at elevated CO₂ and legume N:P was slightly increased. We attribute the less reduced N concentration in legumes at elevated CO₂ to the fact that virtually all legume N originated from symbiotic N₂ fixation (%N_dfa ≈ 90%), and thus legume growth was not limited by soil N. While total plant N was not affected by elevated CO₂, microbial N pools increased by +18% under CO₂ enrichment (P = 0.04) and plant available soil N decreased. Hence, there was a net increase in the overall biotic N pool, largely due increases in the microbial N pool. In order to assess the effects of legumes for ecosystem CO₂ responses and to estimate the degree to which plant growth was P-limited, two greenhouse experiments were conducted, using firstly undisturbed grassland monoliths from the field site, and secondly designed `microcosm' communities on natural soil. Half the microcosms were planted with legumes and half were planted without. Both monoliths and microcosms were exposed to elevated CO₂ and P fertilization in a factored design. After two seasons, plant N pools in both unfertilized monoliths and microcosm communities were unaffected by CO₂ enrichment, similar to what was found in the field. However, when P was added total plant N pools increased at elevated CO₂. This community-level effect originated almost solely from legume stimulation. The results suggest a complex interaction between atmospheric CO₂ concentrations, N and P supply. Overall ecosystem productivity is N-limited, whereas CO₂ effects on legume growth and their N₂ fixation are limited by P. Received: 12 July 1997 / Accepted: 15 April 1998     

Soil moisture dynamics of calcareous grassland under elevated CO2

Water relations of nutrient-poor calcareous grassland under long-term CO₂ enrichment were investigated. Understanding CO₂ effects on soil moisture is critical because productivity in these grasslands is water limited. In general, leaf conductance was reduced at elevated CO₂, but responses strongly depended on date and species. Evapotranspiration (measured as H₂O gas exchange) revealed only small, non-significant reductions at elevated CO₂, indicating that leaf conductance effects were strongly buffered by leaf boundary layer and canopy conductance (leaf area index was not or only marginally increased under elevated CO₂). However, these minute and non-significant responses of water vapour loss accumulated over time and resulted in significantly higher soil moisture in CO₂-enriched plots (gravimetric spot measurements and continuous readings using a network of time-domain reflectometry probes). Differences strongly depended on date, with the smallest effects when soil moisture was very high (after heavy precipitation) and effects were largest at intermediate soil moisture. Elevated CO₂ also affected diurnal soil moisture courses and rewetting of soils after precipitation. We conclude that ecosystem-level controls of the water balance (including soil feedbacks) overshadow by far the physiological effects observed at the leaf level. Indirect effects of CO₂ enrichment mediated by trends in soil moisture will have far-ranging consequences on plant species composition, soil bacterial and faunal activity as well as on soil physical structure and may indirectly also affect hydrology and trace gas emissions and atmospheric chemistry. Received: 21 December 1997 / Accepted: 3 August 1998     

Effects of the frequency and the rate of N enrichment on community structure in a temperate grassland

Aims Nitrogen (N) enrichment caused by human activities threatens biodiversity and alters plant community composition and structure. It has been found that heavy and infrequent N inputs may over-estimate species extinction, but it remains unclear whether plant community structure will equally respond to frequent reactive N enriched conditions.     

Activity of surface-casting earthworms in a calcareous grassland under elevated atmospheric CO2

Earthworms make up the dominant fraction of the biomass of soil animals in most temperate grasslands and have important effects on the structure and function of these ecosystems. We hypothesized that the effects of elevated atmospheric CO₂ on soil moisture and plant biomass production would increase earthworm activity, expressed as surface cast production. Using a screen-aided CO₂ control facility (open top and open bottom rings), eight 1.2-m² grassland plots in Switzerland have been maintained since March 1994 at ambient CO₂ concentrations (350 μl CO₂ l⁻¹) and eight at elevated CO₂ (610 μl CO₂ l⁻¹). Cumulative earthworm surface cast production measured 40 times over 1 year (April 1995–April 1996) in plots treated with elevated CO₂ (2206 g dry mass m⁻² year⁻¹) was 35% greater (P<0.05) than that measured in plant communities maintained at ambient CO₂ (1633 g dry mass m⁻² year⁻¹). At these rates of surface cast production, worms would require about 100 years to egest the equivalent of the amount of soil now found in the Ah horizon (top 15 cm) under current ambient CO₂ concentrations, and 75 years under elevated CO₂. Elevated atmospheric CO₂ had no influence on the seasonality of earthworm activity. Cumulative surface cast production measured over the 7-week period immediately following the 6-week summer dry period in 1995 (no surface casting) was positively correlated (P<0.05) with the mean soil water content calculated over this dry and subsequent wetter period, when viewed across all treatments. However, no correlations were observed with soil temperature or with annual aboveground plant biomass productivity. No CO₂-related differences were observed in total nitrogen (N_tot) and organic carbon (C_org) concentration of surface casts, although concentrations of both elements varied seasonally. The CO₂-induced increase in earthworm surface casting activity corresponded to a 30% increase of the amount of N_tot (8.9 mg N m⁻² vs. 6.9 mg N m⁻²) and C_org (126 mg C m⁻² vs. 94 mg C m⁻²) egested by the worms in one year. Thus, our results demonstrate an important indirect stimulatory effect of elevated atmospheric CO₂ on earthworm activity which may have profound effects on ecosystem function and plant community structure in the long term. Received: 3 November 1996 / Accepted: 11 January 1997     

放牧和刈割对大针茅草原土壤微生物群落结构及多样性的影响

放牧和刈割是内蒙古草原的两种主要利用方式,然而,长期放牧和刈割对大针茅草原土壤微生物群落的影响知之甚少,因此,以内蒙古大针茅草原为研究对象,设置放牧和刈割两种利用方式,以围封不利用为对照,基于高通量测序技术,研究大针茅草原在不同利用方式下土壤微生物组成及多样性的变化,并结合土壤理化因子进一步探究土壤微生物群落组成的主要影响因素。研究结果表明：不同利用方式下土壤细菌α多样性指数无显著差异,而刈割显著提高了土壤真菌Observed＿species、Chao1和ACE指数;土壤细菌群落的优势菌门是变形菌门(Proteobacteria)和放线菌门(Actinobacteria),土壤真菌群落的优势菌门是子囊菌门(Ascomycota)和被孢霉门(Mortierellomycota),不同利用方式下部分微生物类群的相对丰度差异显著,放牧显著提高了细菌群落的变形菌门、疣微菌门(Verrucomicrobia)和芽单胞菌门(Gemmatimonadetes)的相对丰度,刈割显著提高了真菌群落的担子菌门(Basidiomycota)相对丰度,此外,放牧和刈割均显著降低了厚壁菌门(Firmicutes...     

开放式空气二氧化碳浓度增高（FACE）对稻麦轮作土壤微生物数量的影响

利用无锡市安镇的FACE研究平台,在当地正常的栽培及水肥管理条件下,研究了CO2浓度升高对稻麦轮作0-5cm和5～10cm土层土壤微生物数量的影响．结果表明,在水稻拔节期、小麦越冬期与成熟期的0～5cm和5～10cm土层中,FACE处理能显著增加土壤细菌的数量．CO2浓度升高对土壤真菌数量的影响,只有在水稻成熟期0-5cm土层达显著水平,其余均不显著．无论稻季和麦季土壤细菌的数量都远远高于真菌．     

Mycorrhizal fungi reduce the negative effects of nitrogen enrichment on plant community structure in dune grassland

Nitrogen (N) inputs to ecosystems have increased worldwide, often leading to large changes in plant community structure and reducing plant diversity. Yet, the interaction of increased N availability with other factors that determine plant community composition, are still poorly understood. Here, we test whether the impact of N addition on plant communities depends on the presence of arbuscular mycorrhizal fungi (AMF). AMF are widespread plant symbionts that facilitate growth of many plant species. We hypothesize that AM fungi reduce the negative impact of N addition on plant communities by supporting growth of species that are sensitive to N enrichment.We established experimental grassland microcosms consisting of 18 plant species. These microcosms were subjected to high and low N supply and were inoculated with AMF or remained nonmycorrhizal. Both N addition and AMF had a big impact on plant community composition, but with opposite effects. N addition induced a 2.8‐fold increase in grass biomass and reduced legume biomass. Grasses dominated the microcosms at high N supply, especially when AMF were absent. In contrast, AMF enhanced biomass of all legumes species (on average 6.8‐fold) and reduced the relative abundance of grasses. The proportion of legume biomass out of total shoot biomass at high N supply was 19% with AMF and only 3% without AMF. Our results show that responses of plant communities to N enrichment depend on AMF and that AMF can reduce the negative impact of increased N availability on plant community structure by reducing grass dominance.     

Warming and free-air CO2 enrichment alter demographics in four co-occurring grassland species

Species differ in their responses to global changes such as rising CO(2) and temperature, meaning that global changes are likely to change the structure of plant communities. Such alterations in community composition must be underlain by changes in the population dynamics of component species. Here, the impact of elevated CO(2) (550 micromol mol(-1)) and warming (+2 degrees C) on the population growth of four plant species important in Australian temperate grasslands is reported. Data collected from the Tasmanian free-air CO(2) enrichment (TasFACE) experiment between 2003 and 2006 were analysed using population matrix models. Population growth of Themeda triandra, a perennial C(4) grass, was largely unaffected by either factor but population growth of Austrodanthonia caespitosa, a perennial C(3) grass, was reduced substantially in elevated CO(2) plots. Warming and elevated CO(2) had antagonistic effects on population growth of two invasive weeds, Hypochaeris radicata and Leontodon taraxacoides, with warming causing population decline. Analysis of life cycle stages showed that seed production, seedling emergence and establishment were important factors in the responses of the species to global changes. These results show that the demographic approach is very useful in understanding the variable responses of plants to global changes and in elucidating the life cycle stages that are most responsive.     

Changes in the microbial community structure of bacteria,archaea and fungi in response to elevated CO2 and warming in an Australian native grassland soil

The microbial community structure of bacteria, archaea and fungi is described in an Australian native grassland soil after more than 5 years exposure to different atmospheric CO₂ concentrations ([CO₂]) (ambient, + 550 ppm) and temperatures (ambient, + 2°C) under different plant functional types (C ₃ and C ₄ grasses) and at two soil depths (0–5 cm and 5–10 cm). Archaeal community diversity was influenced by elevated [CO₂], while under warming archaeal 16S rRNA gene copy numbers increased for C ₄ plant Themeda triandra and decreased for the C ₃ plant community (P < 0.05). Fungal community diversity resulted in three groups based upon elevated [CO₂], elevated [CO₂] plus warming and ambient [CO₂]. Overall bacterial community diversity was influenced primarily by depth. Specific bacterial taxa changed in richness and relative abundance in response to climate change factors when assessed by a high‐resolution 16S rRNA microarray (PhyloChip). Operational taxonomic unit signal intensities increased under elevated [CO₂] for both Firmicutes and Bacteroidetes, and increased under warming for Actinobacteria and Alphaproteobacteria. For the interaction of elevated [CO₂] and warming there were 103 significant operational taxonomic units (P < 0.01) representing 15 phyla and 30 classes. The majority of these operational taxonomic units increased in abundance for elevated [CO₂] plus warming plots, while abundance declined in warmed or elevated [CO₂] plots. Bacterial abundance (16S rRNA gene copy number) was significantly different for the interaction of elevated [CO₂] and depth (P < 0.05) with decreased abundance under elevated [CO₂] at 5–10 cm, and for Firmicutes under elevated [CO₂] (P < 0.05). Bacteria, archaea and fungi in soil responded differently to elevated [CO₂], warming and their interaction. Taxa identified as significantly climate‐responsive could show differing trends in the direction of response (‘+’ or ‘?’) under elevated CO₂ or warming, which could then not be used to predict their interactive effects supporting the need to investigate interactive effects for climate change. The approach of focusing on specific taxonomic groups provides greater potential for understanding complex microbial community changes in ecosystems under climate change.     

Changes in soil microbial community structure and function following degradation in a temperate grassland

温带草原退化后土壤微生物群落结构和功能的变化 草原退化是草原生产力维持面临的一个重大挑战,这一过程显著影响着草原生态系统的能量流动和土壤养分变化过程,进而直接或间接地影响着土壤微生物。我们的研究目的首先是调查不同草原退化程度(即未退化、中度退化和严重退化)如何影响着内蒙古温带草原的土壤微生物组成、多样性和功能,其次是阐明哪些生物和非生物因素导致了这些变化。我们的研究主要通过高通量测序技术分析土壤微生物的群落组成,并且采用FAPROTAX工具和FUNGuild工具分别预测细菌群落和真菌群落的功能。研究发现：草原退化显著降低了土壤细菌的多样性,但对真菌多样性影响不大。地下生物量、土壤有机碳和总氮与细菌的多样性变化呈显著正相关关系。草原退化显著增加了绿弯菌门的相对丰度(由2.48%提高到8.40%),降低了厚壁菌门的相对丰度(由3.62%降低到1.08%)。其次,草原退化也显著增加了球囊菌门的相对丰度(从0.17%提高到1.53%),降低了担子菌门的相对丰度(从19.30%降低到4.83%)。致病菌的相对丰度在草原退化过程中显著下降。此外,草原退化对土壤细菌群落的功能有显著的影响,尤其是与土壤碳氮循环相关的土壤细菌群落。我们的结果表明,土壤细菌群落对草原退化的响应比真菌群落更敏感。     

氯嘧磺隆对土壤微生物群落结构的影响

采用磷脂脂肪酸(PLFA)法研究了黑龙江省苇河地区不同氯嘧磺隆施药历史下土壤微生物群落结构的差异, 并测定了土壤中氯嘧磺隆的残留量.结果表明：不同施药历史下氯嘧磺隆在土壤中的残留量均很低;随着施用氯嘧磺隆年限的增加,土壤微生物的PLFA总量减少, 革兰氏阴性菌/革兰氏阳性菌（GN/GP）和真菌/细菌比降低,土壤微生物的压力指数增加.主成分分析表明,氯嘧磺隆显著改变了大豆田土壤微生物群落结构.     

Plant community dynamics in a calcareous grassland under climate change manipulations

This study investigates the effects of field manipulations of local climate to determine the potential impact of climate change on plant community dynamics in a calcareous grassland. The experimental site is located in a grassland at the Wytham estate, Oxfordshire, UK. The one hectare study area is within a 10 ha abandoned arable field on Jurassic corallian limestone. Two climate change scenarios were used: warmer winters with increased summer rainfall and warmer winters with summer drought. Plant cover and species richness were significantly increased in plots receiving supplemented summer rainfall, while the amount of litter was significantly reduced. Litter formation was significantly increased by winter warming and drought. The responses of the plant community to the climate manipulations were related to the life-history attributes of the dominant species. Seedling recruitment was limited by microsite availability, which also varied in the different climate manipulations. The results are discussed in terms of successional dynamics. They suggest that warmer winters may delay succession, as gap formation in the sward will provide sites for colonisation of annuals, thereby enabling their persistence in the sward. Under wetter conditions during summer, perennial grasses tend to close the sward, thereby inhibiting the establishment of later successional species.