Plant species, atmospheric CO2 and soil N interactively or additively control C allocation within plant-soil systems

Two plant species, Medicago truncatula (legume) and Avena sativa (non-legume), were grown in low-or high-N soils under two CO₂ concentrations to test the hypothesis whether C allocation within plant-soil system is interactively or additively controlled by soil N and atmospheric CO₂ is dependent upon plant species. The results showed the interaction between plant species and soil N had a significant impact on microbial activity and plant growth. The interaction between CO₂ and soil N had a significant impact on soil soluble C and soil microbial biomass C under Madicago but not under Avena. Although both CO₂ and soil N affected plant growth significantly, there was no interaction between CO₂ and soil N on plant growth. In other words, the effects of CO₂ and soil N on plant growth were additive. We considered that the interaction between N₂ fixation trait of legume plant and elevated CO₂ might have obscured the interaction between soil N and elevated CO₂ on the growth of legume plant. In low-N soil, the shoot-to-root ratio of Avena dropped from 2.63±0.20 in the early growth stage to 1.47±0.03 in the late growth stage, indicating that Avena plant allocated more energy to roots to optimize nutrient uptake (i.e. N) when soil N was limiting. In high-N soil, the shoot-to-root ratio of Medicago increased significantly over time (from 2.45±0.30 to 5.43±0.10), suggesting that Medicago plants allocated more energy to shoots to optimize photosynthesis when N was not limiting. The shoot-to-root ratios were not significantly different between two CO₂ levels.     

Plant species, atmospheric CO2 and soil N interactively or additively control C allocation within plant-soil systems

Two plant species, Medicago truncatula (legume) and Avena sativa (non-legume), were grown in low-or high-N soils under two CO₂ concentrations to test the hypothesis whether C allocation within plant-soil system is interactively or additively controlled by soil N and atmospheric CO₂ is dependent upon plant species. The results showed the interaction between plant species and soil N had a significant impact on microbial activity and plant growth. The interaction between CO₂ and soil N had a significant impact on soil soluble C and soil microbial biomass C under Madicago but not under Avena. Although both CO₂ and soil N affected plant growth significantly, there was no interaction between CO₂ and soil N on plant growth. In other words, the effects of CO₂ and soil N on plant growth were additive. We considered that the interaction between N₂ fixation trait of legume plant and elevated CO₂ might have obscured the interaction between soil N and elevated CO₂ on the growth of legume plant. In low-N soil, the shoot-to-root ratio of Avena dropped from 2.63±0.20 in the early growth stage to 1.47±0.03 in the late growth stage, indicating that Avena plant allocated more energy to roots to optimize nutrient uptake (i.e. N) when soil N was limiting. In high-N soil, the shoot-to-root ratio of Medicago increased significantly over time (from 2.45±0.30 to 5.43±0.10), suggesting that Medicago plants allocated more energy to shoots to optimize photosynthesis when N was not limiting. The shoot-to-root ratios were not significantly different between two CO₂ levels.     

Carbon and nitrogen allocation in Lolium perenne in response to elevated atmospheric CO2 with emphasis on soil carbon dynamics

The effect of elevated CO2 on the carbon and nitrogen distribution within perennial ryegrass (L. perenne L.) and its influence on belowground processes were investigated. Plants were homogeneously 14C-labelled in two ESPAS growth chambers in a continuous 14C-CO2 atmosphere of 350 and 700 L L-1 CO2 and at two soil nitrogen regimes, in order to follow the carbon flow through all plant and soil compartments.After 79 days, elevated CO2 increased the total carbon uptake by 41 and 21% at low (LN) and high nitrogen (HN) fertilisation, respectively. Shoot growth remained unaffected, whereas CO2 enrichment stimulated root growth by 46% and the root/soil respiration by 111%, irrespective of the nitrogen concentration. The total 14C-soil content increased by 101 and 28% at LN and HN, respectively. The decomposition of the native soil organic matter was not affected either by CO2 or by the nitrogen treatment.Elevated CO2 did not change the total nitrogen uptake of the plant either at LN or at HN. Both at LN and HN elevated CO2 significantly increased the total amount of nitrogen taken up by the roots and decreased the absolute and relative amounts translocated to the shoots.The amount of soil nitrogen immobilised by micro-organisms and the size of the soil microbial biomass were not affected by elevated CO2, whereas both were significantly increased at the higher soil N content.Most striking was the 88% increase in net carbon input into the soil expressed as: 14C-roots plus total 14C-soil content minus the 12C-carbon released by decomposition of native soil organic matter. The net carbon input into the soil at ambient CO2 corresponded with 841 and 1662 kg ha-1 at LN and HN, respectively. Elevated CO2 increased these amounts with an extra carbon input of 950 and 1056 kg ha-1. Combined with a reduced decomposition rate of plant material grown at elevated CO2 this will probably lead to carbon storage in grassland soils resulting in a negative feed back on the increasing CO2 concentration of the atmosphere.     

Influence of Doubled CO2 on Plant Growth and Soil Microbial Biomass C and N

Salix babylonica L., Triticum aestivum L., Chenopodium album L. and Amaranthus cruemus L. were grown in the N-deficient soil in open-top chambers blown with ambient or doubled ambient CO2 air, and their growth was measured. Soil samples were collected to assess the influence of doubled CO2 on the soil microbial biomass C (Cmic) and N (Nmic). Results showed that the biomass of shoot and root was increased by doubled CO2 in the four species of plants. Doubled CO2 increased Cmic in S. babylonica and decreased Cmic in T. aestivum and C. album. On the other hand, Nmic in three species except T. aestivum was stimulated by doubled CO2. Doubled CO2 had no significant effect on Cmic in A. cruentus and Nmic in T. aestivum. However, the ratios of Cmic- to -Nmic of all four species were consistently declined under doubled CO2 treatment. It implies that CO2 enrichment may have positive influence on the quality of organic matter of N-low soil in global change.     

全球大气CO2浓度升高对土壤微生物的影响

全球大气CO2浓度升高对土壤微生物生态系统的影响已引起广泛关注。本文从土壤微生物群落结构、微生物区系、土壤呼吸、微生物生物量以及土壤酶活性方面对大气高浓度CO2的响应进行了综述。由于提供高浓度CO2的实验系统、所选植物材料以及土壤特性等的不同,大气CO2浓度升高对土壤微生物群落结构、微生物区系、土壤呼吸、微生物生物量以及土壤酶活性的影响并未得出一致结论。但高浓度CO2对土壤微生物生态系统的影响是存在的。     

Legumes regulate grassland soil N cycling and its response to variation in species diversity and N supply but not CO2

Legumes are an important component of plant diversity that modulate nitrogen (N) cycling in many terrestrial ecosystems. Limited knowledge of legume effects on soil N cycling and its response to global change factors and plant diversity hinders a general understanding of whether and how legumes broadly regulate the response of soil N availability to those factors. In a 17‐year study of perennial grassland species grown under ambient and elevated (+180 ppm) CO₂ and ambient and enriched (+4 g N m^?2 year^?1) N environments, we compared pure legume plots with plots dominated by or including other herbaceous functional groups (and containing one or four species) to assess the effect of legumes on N cycling (net N mineralization rate and inorganic N pools). We also examined the effects of numbers of legume species (from zero to four) in four‐species mixed plots on soil N cycling. We hypothesized that legumes would increase N mineralization rates most in those treatments with the greatest diversity and the greatest relative limitation by and competition for N. Results partially supported these hypotheses. Plots with greater dominance by legumes had greater soil nitrate concentrations and mineralization rates. Higher species richness significantly increased the impact of legumes on soil N metrics, with 349% and 505% higher mineralization rates and nitrate concentrations in four‐species plots containing legumes compared to legume‐free four‐species plots, in contrast to 185% and 129% greater values, respectively, in pure legume than nonlegume monoculture plots. N‐fertilized plots had greater legume effects on soil nitrate, but lower legume effects on net N mineralization. In contrast, neither elevated CO₂ nor its interaction with legumes affected net N mineralization. These results indicate that legumes markedly influence the response of soil N cycling to some, but not all, global change drivers.     

Response of soil biota to elevated atmospheric CO2 in poplar model systems

We tested the hypotheses that increased belowground allocation of carbon by hybrid poplar saplings grown under elevated atmospheric CO₂ would increase mass or turnover of soil biota in bulk but not in rhizosphere soil. Hybrid poplar saplings (Populus×euramericana cv. Eugenei) were grown for 5 months in open-bottom root boxes at the University of Michigan Biological Station in northern, lower Michigan. The experimental design was a randomized-block design with factorial combinations of high or low soil N and ambient (34 Pa) or elevated (69 Pa) CO₂ in five blocks. Rhizosphere microbial biomass carbon was 1.7 times greater in high-than in low-N soil, and did not respond to elevated CO₂. The density of protozoa did not respond to soil N but increased marginally (P < 0.06) under elevated CO₂. Only in high-N soil did arbuscular mycorrhizal fungi and microarthropods respond to CO₂. In high-N soil, arbuscular mycorrhizal root mass was twice as great, and extramatrical hyphae were 11% longer in elevated than in ambient CO₂ treatments. Microarthropod density and activity were determined in situ using minirhizotrons. Microarthropod density did not change in response to elevated CO₂, but in high-N soil, microarthropods were more strongly associated with fine roots under elevated than ambient treatments. Overall, in contrast to the hypotheses, the strongest response to elevated atmospheric CO₂ was in the rhizosphere where (1) unchanged microbial biomass and greater numbers of protozoa (P < 0.06) suggested faster bacterial turnover, (2) arbuscular mycorrhizal root length increased, and (3) the number of microarthropods observed on fine roots rose. Received: 18 March 1997 / Accepted: 5 August 1997     

Effects of elevated atmospheric CO2 on soil microbiota in calcareous grassland

Microbial responses to three years of CO₂ enrichment (600 μL L^–1) in the field were investigated in calcareous grassland. Microbial biomass carbon (C) and soil organic C and nitrogen (N) were not significantly influenced by elevated CO₂. Microbial C:N ratios significantly decreased under elevated CO₂ (– 15%, P = 0.01) and microbial N increased by + 18% (P = 0.04). Soil basal respiration was significantly increased on one out of 7 sampling dates (+ 14%, P = 0.03; December of the third year of treatment), whereas the metabolic quotient for CO₂ (qCO₂ = basal respiration/microbial C) did not exhibit any significant differences between CO₂ treatments. Also no responses of microbial activity and biomass were found in a complementary greenhouse study where intact grassland turfs taken from the field site were factorially treated with elevated CO₂ and phosphorus (P) fertilizer (1 g P m^–2 y^–1). Previously reported C balance calculations showed that in the ecosystem investigated growing season soil C inputs were strongly enhanced under elevated CO₂. It is hypothesized that the absence of microbial responses to these enhanced soil C fluxes originated from mineral nutrient limitations of microbial processes. Laboratory incubations showed that short-term microbial growth (one week) was strongly limited by N availability, whereas P was not limiting in this soil. The absence of large effects of elevated CO₂ on microbial activity or biomass in such nutrient-poor natural ecosystems is in marked contrast to previously published large and short-term microbial responses to CO₂ enrichment which were found in fertilized or disturbed systems. It is speculated that the absence of such responses in undisturbed natural ecosystems in which mineral nutrient cycles have equilibrated over longer periods of time is caused by mineral nutrient limitations which are ineffective in disturbed or fertilized systems and that therefore microbial responses to elevated CO₂ must be studied in natural, undisturbed systems.     

Elevated atmospheric CO2 increases microbial growth rates in soil: results of three CO2 enrichment experiments

Increasing the belowground translocation of assimilated carbon by plants grown under elevated CO₂ can cause a shift in the structure and activity of the microbial community responsible for the turnover of organic matter in soil. We investigated the long‐term effect of elevated CO₂ in the atmosphere on microbial biomass and specific growth rates in root‐free and rhizosphere soil. The experiments were conducted under two free air carbon dioxide enrichment (FACE) systems: in Hohenheim and Braunschweig, as well as in the intensively managed forest mesocosm of the Biosphere 2 Laboratory (B2L) in Oracle, AZ. Specific microbial growth rates (μ) were determined using the substrate‐induced respiration response after glucose and/or yeast extract addition to the soil. For B2L and both FACE systems, up to 58% higher μ were observed under elevated vs. ambient CO₂, depending on site, plant species and N fertilization. The μ‐values increased linearly with atmospheric CO₂ concentration at all three sites. The effect of elevated CO₂ on rhizosphere microorganisms was plant dependent and increased for: Brassica napus=Triticum aestivum<Beta vulgaris<Populus deltoides. N deficiency affected microbial growth rates directly (N limitation) and indirectly (changing the quantity of fine roots). So, 50% decrease in N fertilization caused the overall increase or decrease of microbial growth rates depending on plant species. The μ‐value increase was lower for microorganisms growing on yeast extract then for those growing on glucose, i.e. the effect of elevated CO₂ was smoothed on rich vs. simple substrate. So, the r/K strategies ratio can be better revealed by studying growth on simple (glucose) than on rich substrate mixtures (yeast extract). Our results clearly showed that the functional characteristics of the soil microbial community (i.e. specific growth rates) rather than total microbial biomass amount are sensitive to increased atmospheric CO₂. We conclude that the more abundant available organics released by roots at elevated CO₂ altered the ecological strategy of the soil microbial community specifically a shift to a higher contribution of fast‐growing r‐selected species was observed. These changes in functional structure of the soil microbial community may counterbalance higher C input into the soil under elevated atmospheric CO₂ concentration.     

Plant and microbial N acquisition under elevated atmospheric CO2 in two mesocosm experiments with annual grasses

The impact of elevated CO₂ on terrestrial ecosystem C balance, both in sign or magnitude, is not clear because the resulting alterations in C input, plant nutrient demand and water use efficiency often have contrasting impacts on microbial decomposition processes. One major source of uncertainty stems from the impact of elevated CO₂ on N availability to plants and microbes. We examined the effects of atmospheric CO₂ enrichment (ambient+370 μmol mol^?1) on plant and microbial N acquisition in two different mesocosm experiments, using model plant species of annual grasses of Avena barbata and A. fatua, respectively. The A. barbata experiment was conducted in a N‐poor sandy loam and the A. fatua experiment was on a N‐rich clayey loam. Plant–microbial N partitioning was examined through determining the distribution of a ¹⁵N tracer. In the A. barbata experiment, ¹⁵N tracer was introduced to a field labeling experiment in the previous year so that ¹⁵N predominantly existed in nonextractable soil pools. In the A. fatua experiment, ¹⁵N was introduced in a mineral solution [(¹⁵NH₄)₂SO₄ solution] during the growing season of A. fatua. Results of both N budget and ¹⁵N tracer analyses indicated that elevated CO₂ increased plant N acquisition from the soil. In the A. barbata experiment, elevated CO₂ increased plant biomass N by ca. 10% but there was no corresponding decrease in soil extractable N, suggesting that plants might have obtained N from the nonextractable organic N pool because of enhanced microbial activity. In the A. fatua experiment, however, the CO₂‐led increase in plant biomass N was statistically equal to the reduction in soil extractable N. Although atmospheric CO₂ enrichment enhanced microbial biomass C under A. barbata or microbial activity (respiration) under A. fatua, it had no significant effect on microbial biomass N in either experiment. Elevated CO₂ increased the colonization of A. fatua roots by arbuscular mycorrhizal fungi, which coincided with the enhancement of plant competitiveness for soluble soil N. Together, these results suggest that elevated CO₂ may tighten N cycling through facilitating plant N acquisition. However, it is unknown to what degree results from these short‐term microcosm experiments can be extrapolated to field conditions. Long‐term studies in less‐disturbed soils are needed to determine whether CO₂‐enhancement of plant N acquisition can significantly relieve N limitation over plant growth in an elevated CO₂ environment.     

大气CO2浓度升高对不同施氮土壤酶活性的影响

利用中国唯一的无锡FACE（Free-air CO2 enrichment,开放式空气CO2浓度升高）平台,研究了大气CO2浓度升高对土壤β-葡糖苷酶、转化酶、脲酶、酸性磷酸酶、-氨基葡糖苷酶的影响。研究发现,不同氮肥处理下大气CO2浓度升高对某些土壤酶活性的影响不同。在低氮施肥处理中,大气CO2浓度升高显著降低-葡糖苷酶活性,但是在高氮施肥处理下,大气CO2浓度升高显著增加β-葡糖苷酶活性。在低氮和常氮施肥处理中大气CO2浓度升高显著增加了土壤脲酶活性,但在高氮水平下影响不显著。在低氮、常氮施肥处理中,大气CO2浓度升高对土壤酸性磷酸酶活性没有影响,而在高氮施肥处理中显著增强了土壤中磷酸酶活性。大气CO2浓度升高对土壤转化酶活性和-氨基葡糖苷酶的活性有增加趋势,但影响不显著。研究还发现,在不同的CO2浓度下,土壤酶活性对不同氮肥处理的响应也不同。在正常CO2浓度下,土壤中β-葡糖苷酶活性随着氮肥施用量的增加而降低,而在大气CO2浓度升高条件下,却随着氮肥施用量的增加而增加。在大气CO2浓度升高条件下,高氮施肥显著增加了转化酶和酸性磷酸酶活性,而在正常CO2浓度下,影响不显著。在大气CO2浓度升高条件下,氮肥处理对脲酶活性的影响不大,但在正常CO2浓度下,脲酶活性随着氮肥施用量的增加而增加。氮肥对β-氨基葡糖苷酶活性的影响不明显。     

CO_2倍增对植物生长和土壤微生物生物量碳、氮的影响

关于大气ＣＯ２浓度倍增（即为７００μｍｏｌＣＯ２·ｍｏｌ－１空气）将对植物生长产生诸多影响,已有大量报道［１,２］。但ＣＯ２倍增对植物及所在土壤中微生物影响的研究甚少［３,４］。土壤微生物是陆地生态系统中最活跃的成分,担负着分解动植物残体的重要作用,...     

Divergent terrestrial responses of soil N2O emissions to different levels of elevated CO2 and temperature

Understanding the responses of soil nitrous oxide (N₂O) emissions from terrestrial ecosystems to future CO₂ enrichment and warming is critical for the development of mitigation and adaptation policies. The effects of continuous increase in elevated CO₂ (EC) and elevated temperature (ET) on N₂O emissions are not fully known. We synthesized 209 measurements from 70 published studies and carried out a meta-analysis to examine individual and interactive effects of EC and ET on N₂O emissions from grasslands, croplands and forests. On average, a significant increase of 23% in N₂O emissions was observed under EC across all case studies. EC did not affect N₂O emissions from grasslands or forests, but significantly increased N₂O emissions in croplands by 38%. The extent of ET effects on N₂O emissions was nonsignificant and there was no significant difference in N₂O emission responses among these three terrestrial systems. ET only promoted N₂O emissions in forest by about 32% when ET was less than 2°C. The interactive effect of EC and ET on N₂O emissions was significantly synergistic, showing a greater increase than the sum of the effects caused by EC and ET alone. Our findings indicated that the combination of EC and ET substantially promoted soil N₂O and highlighted the urgent need to explore its mechanisms to better understand N₂O responses under future climate change.     

Plant growth chambers for the simultaneous control of soil and air temperatures, and of atmospheric carbon dioxide concentration

Many facilities for growing plants at elevated atmospheric concentrations of CO₂ ([CO₂]) neglect the control of temperature, especially of the soil. Soil and root temperatures in conventional, free-standing pots often exceed those which would occur in the field at a given air temperature. A plant growth facility is described in which atmospheric CO₂ can be maintained at different concentrations while soil and air temperatures mimic spatial and temporal patterns seen in the field. It consists of glasshouse-located chambers in which [CO₂] is monitored by an infra-red gas analyser and maintained by injection of CO₂ from a cylinder. Air is cooled by a heat exchange unit. Plants grow in soil in 1.2 m long containers that are surrounded by cooling coils and thermal insulation. Both [CO₂] and temperature are controlled by customized software. Air temperature is programmed to follow a sine function of diurnal time. Soil temperature at a depth of 0.55 m is programmed to be constant. Temperature at 0.1 m depth varies as a damped, lagged function of air temperature; that at 1.0 m as a similar function of the 0.55 m temperature. [CO₂] is maintained within 20 μmol mol^?1 of target concentrations during daylight. A feature of the system is that plant material is labelled with a ¹³C enrichment different from that of carbon in soil organic matter. The operation of the system is illustrated with data collected in an experiment with spring wheat (Triticum aestivum L., cv Tonic) grown at ambient [CO₂] and at [CO₂] 350 μmol mol^?1 greater than ambient.     

大气CO2浓度升高对红桦幼苗根系的影响

应用封闭式生长室系统,研究了CO2浓度升高对红桦(Betula albosinensis)幼苗的根/冠、粗根和细根的干质量、非结构性碳水化合物类含量、碳含量和碳/氮、氮和磷的含量及氮磷吸收量的影响。结果表明:CO2浓度升高使红桦幼苗粗根和细根的干质量增加,同时根/冠值显著升高,表明CO2浓度升高使红桦幼苗生物量向根系的分配增加;与对照相比,粗根的还原糖、蔗糖和总可溶性糖含量显著增加,而在细根中没有显著变化;粗根、细根的淀粉和总的非结构性碳水化合物含量显著增加;CO2浓度升高下粗根和细根的碳含量有升高的趋势但未达到显著水平,同时氮含量降低,碳/氮值升高;氮的吸收量在粗根和细根中均无显著变化。上述结果表明,CO2浓度升高下红桦幼苗根系氮含量下降是由非结构性碳水化合物(主要是淀粉)含量升高和(或)根系生物量增加产生的稀释效应引起的。     

大气二氧化碳浓度与温度升高对红桦幼苗养分积累和分配的影响

利用封闭式生长室,研究了CO2浓度升高(环境CO2 350 μmol·mol-1,EC)、温度升高(环境温度 2 ℃,ET)以及二者同时升高(ECT)对川西亚高山红桦幼苗养分积累和分配的影响.结果表明:经过一个生长季, EC处理下红桦幼苗单株N、P、K积累比对照分别增加44%、45%和11%(P《0.05),ET处理下分别增加37%、76%和9%(P《0.05),ECT处理下分别增加24%、88%和20% (P《0.05).EC处理使N向红桦幼苗叶中分配的比例降低11.68%(P《0.05),向枝、茎、根中分配的比例分别增加2.95%、3.39%和5.34%(P》0.05);ET处理使N向叶中分配的比例增加11.09%(P《0.05),向枝、茎、根中分配的比例分别降低0.69%、10.35%和0.05%(P》0.05).ECT处理下N的分配格局与EC处理相似.3种处理下P和K在红桦幼苗中的分配变化差异较大,CO2浓度和温度升高可能促进植物养分的积累,改变养分在植物各器官间的分配.     

大气CO2浓度升高对植物根系的影响

植物长期生长在CO2浓度不断升高的环境中,其结构和功能都将受到影响,这种影响不仅表现在植物的地上部分,同时也表现在植物的地下部分(根系),尤其是细根的长度、直径、产量、周转以及根与枝的分配模式等方面。植物根系结构和功能的改变影响植物地上部分和生态系统物质循环中的碳动态及土壤中碳库的变化。目前有关大气CO2浓度升高对根系动态影响的研究报道主要包括大气CO2浓度升高对根系结构(直径、分枝、长度、数量等)和根系生理(周转率、产量、碳分配模式等)的影响2个方面。目前,该领域研究还存在一些不足,例如在CO2浓度升高条件下,对植物根系内部的调控机制,以及由其引起的物质循环和能量流动的动态变化的了解较少;至今没有令人信服的证据说明大气CO2浓度升高使根系周转升高还是降低。今后应加强研究在CO2浓度升高条件下根系的周转变化和光合产物分配模式变化,CO2浓度升高和外界环境因素的共同作用对根系的影响,以及采用不同研究方法和研究对象在不同立地条件下开展升高CO2浓度对根系影响的对比研究等。     

Contrasting growth response of an N2-fixing and non-fixing forb to elevated CO2: dependence on soil N supply

With the ability to symbiotically fix atmospheric N₂, legumes may lack the N-limitations thought to constrain plant response to elevated concentrations of atmospheric CO₂. The growth and photosynthetic responses of two perennial grassland species were compared to test the hypotheses that (1) the CO₂ response of wild species is limited at low N availability, (2) legumes respond to a greater extent than non-fixing forbs to elevated CO₂, and (3) elevated CO₂ stimulates symbiotic N₂ fixation, resulting in an increased amount of N derived from the atmosphere. This study investigated the effects of atmospheric CO₂ concentration (365 and 700 mol mol^–1) and N addition on whole plant growth and C and N acquisition in an N₂-fixing legume (Lupinus perennis) and a non-fixing forb (Achillea millefolium) in controlled-chamber environments. To evaluate the effects of a wide range of N availability on the CO₂ response, we incorporated six levels of soil N addition starting with native field soil inherently low in N (field soil + 0, 4, 8, 12, 16, or 20 g N m^–2 yr^–1). Whole plant growth, leaf net photosynthetic rates (A), and the proportion of N derived from N₂ fixation were determined in plants grown from seed over one growing season. Both species increased growth with CO₂enrichment, but this response was mediated by N supply only for the non-fixer, Achillea. Its response depended on mineral N supply as growth enhancements under elevated CO₂ increased from 0% in low N soil to +25% at the higher levels of N addition. In contrast, Lupinus plants had 80% greater biomass under elevated CO₂ regardless of N treatment. Although partial photosynthetic acclimation to CO₂ enrichment occurred, both species maintained comparably higher A in elevated compared to ambient CO₂ (+38%). N addition facilitated increased A in Achillea, however, in neither species did additional N availability affect the acclimation response of A to CO₂. Elevated CO₂ increased plant total N yield by 57% in Lupinus but had no effect on Achillea. The increased N in Lupinus came from symbiotic N₂ fixation, which resulted in a 47% greater proportion of N derived from fixation relative to other sources of N. These results suggest that compared to non-fixing forbs, N₂-fixers exhibit positive photosynthetic and growth responses to increased atmospheric CO₂ that are independent of soil N supply. The enhanced amount of N derived from N₂ fixation under elevated CO₂ presumably helps meet the increased N demand in N₂-fixing species. This response may lead to modified roles of N₂-fixers and N₂-fixer/non-fixer species interactions in grassland communities, especially those that are inherently N-poor, under projected rising atmospheric CO₂.     

Elevated atmospheric CO2 alters microbial population structure in a pasture ecosystem

An increase in concentration of atmospheric CO₂ is one major factor influencing global climate change. Among the consequences of such an increase is the stimulation of plant growth and productivity. Below‐ground microbial processes are also likely to be affected indirectly by rising atmospheric CO₂ levels, through increased root growth and rhizodeposition rates. Because changes in microbial community composition might have an impact on symbiotic interactions with plants, the response of root nodule symbionts to elevated atmospheric CO₂ was investigated. In this study we determined the genetic structure of 120 Rhizobium leguminosarum bv. trifolii isolates from white clover plants exposed to ambient (350 μmol mol^?1) or elevated (600 μmol mol^?1) atmospheric CO₂ concentrations in the Swiss FACE (Free‐Air‐Carbon‐Dioxide‐Enrichment) facility. Polymerase Chain Reaction (PCR) fingerprinting of genomic DNA showed that the isolates from plants grown under elevated CO₂ were genetically different from those isolates obtained from plants grown under ambient conditions. Moreover, there was a 17% increase in nodule occupancy under conditions of elevated atmospheric CO₂ when strains of R. leguminosarum bv. trifolii isolated from plots exposed to CO₂ enrichment were evaluated for their ability to compete for nodulation with those strains isolated from ambient conditions. These results indicate that a shift in the community composition of R. leguminosarum bv. trifolii occurred as a result of an increased atmospheric CO₂ concentration, and that elevated atmospheric CO₂ affects the competitive ability of root nodule symbionts, most likely leading to a selection of these particular strains to nodulate white clover.