Ten years of free-air CO2 enrichment altered the mobilization of N from soil in Lolium perenne L. swards

Effects of free‐air carbon dioxide enrichment (FACE, 60 Pa pCO₂) on plant growth as compared with ambient pCO₂ (36 Pa) were studied in swards of Lolium perenne L. (perennial ryegrass) at two levels of N fertilization (14 and 56 g m^?2 a^?1) from 1993 to 2002. The objectives were to determine how plant growth responded to the availability of C and N in the long term and how the supply of N to the plant from the two sources of N in the soil, soil organic matter (SOM) and mineral fertilizer, varied over time. In three field experiments, ¹⁵N‐labelled fertilizer was used to distinguish the sources of available N. In 1993, harvestable biomass under elevated pCO₂ was 7% higher than under ambient pCO₂. This relative pCO₂ response increased to 32% in 2002 at high N, but remained low at low N. Between 1993 and 2002, the proportions and amounts of N in harvestable biomass derived from SOM (excluding remobilized fertilizer) were, at high N, increasingly higher at elevated pCO₂ than at ambient pCO₂. Two factorial experiments confirmed that at high N, but not at low N, a higher proportion of N in harvestable biomass was derived from soil (including remobilized fertilizer) following 7 and 9 years of elevated pCO₂, when compared with ambient pCO₂. It is suggested that N availability in the soil initially limited the pCO₂ response of harvestable biomass. At high N, the limitation of plant growth decreased over time as a result of the stimulated mobilization of N from soil, especially from SOM. Consequently, harvestable biomass increasingly responded to elevated pCO₂. The underlying mechanisms which contributed to the increased mobilization of N from SOM under elevated pCO₂ are discussed. This study demonstrated that there are feedback mechanisms in the soil which are only revealed during long‐term field experiments. Such investigations are thus, a prerequisite for understanding the responses of ecosystems to elevated pCO₂ and N supply.     

Comparison of gas use efficiency and treatment uniformity in a forest ecosystem exposed to elevated [CO2] using pure and prediluted free-air CO2 enrichment technology

A direct comparison of treatment uniformity and CO₂ use of pure and prediluted free-air CO₂ enrichment (FACE) systems was conducted in a forest ecosystem. A vertical release pure CO₂ fumigation system was superimposed on an existing prediluted CO₂ fumigation system and operated on alternate days. The FACE system using prediluted CO₂ fumigation technology exhibited less temporal and spatial variability than the pure CO₂ fumigation system. The pure CO₂ fumigation system tended to over-fumigate the upwind portions of the plot and used 25% more CO₂ than the prediluted CO₂ fumigation system. The increased CO₂ use by the pure CO₂ system was exacerbated at low wind speeds. It is not clear if this phenomenon will also be observed in plots with smaller diameters and low-stature vegetation.     

A free-air enrichment system for exposing tall forest vegetation to elevated atmospheric CO2

A free-air CO₂ enrichment (FACE) system was designed to permit the experimental exposure of tall vegetation such as stands of forest trees to elevated atmospheric CO₂ concentrations ([CO₂]_a) without enclosures that alter tree microenvironment. We describe a prototype FACE system currently in operation in forest plots in a maturing loblolly pine (Pinus taeda L.) stand in North Carolina, USA. The system uses feedback control technology to control [CO₂] in a 26 m diameter forest plot that is over 10 m tall, while monitoring the 3D plot volume to characterize the whole-stand CO₂ regime achieved during enrichment. In the second summer season of operation of the FACE system, atmospheric CO₂ enrichment was conducted in the forest during all daylight hours for 96.7% of the scheduled running time from 23 May to 14 October with a preset target [CO₂] of 550 μmol mol^–1, ≈ 200 μmol mol^–1 above ambient [CO₂]. The system provided spatial and temporal control of [CO₂] similar to that reported for open-top chambers over trees, but without enclosing the vegetation. The daily average daytime [CO₂] within the upper forest canopy at the centre of the FACE plot was 552 ± 9 μmol mol^–1 (mean ± SD). The FACE system maintained 1-minute average [CO₂] to within ± 110 μmol mol^–1 of the target [CO₂] for 92% of the operating time. Deviations of [CO₂] outside of this range were short-lived (most lasting < 60 s) and rare, with fewer than 4 excursion events of a minute or longer per day. Acceptable spatial control of [CO₂] by the system was achieved, with over 90% of the entire canopy volume within ± 10% of the target [CO₂] over the exposure season. CO₂ consumption by the FACE system was much higher than for open-top chambers on an absolute basis, but similar to that of open-top chambers and branch bag chambers on a per unit volume basis. CO₂ consumption by the FACE system was strongly related to windspeed, averaging 50 g CO₂ m^–3 h^–1 for the stand for an average windspeed of 1.5 m s^–1 during summer. The [CO₂] control results show that the free-air approach is a tractable way to study long-term and short-term alterations in trace gases, even within entire tall forest ecosystems. The FACE approach permits the study of a wide range of forest stand and ecosystem processes under manipulated [CO₂]_a that were previously impossible or intractable to study in true forest ecosystems.     

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.     

Saprobic microfungi under Lolium perenne and Trifolium repens at different fertilization intensities and elevated atmospheric CO2 concentration

Anthropogenic increases in atmospheric CO₂ concentration and the connected deposition of organic matter into the soil influence the occurrence of decomposers who regulate carbon release back into the atmosphere. The effects of increased concentration of atmospheric carbon dioxide, plant species cover quality and nitrogen (N) fertilization on the coenosis composition of soil saprobic microfungi were studied under field conditions (Swiss Free Air Carbon Dioxide Enrichment experiment). In total, 42 species of microfungi were detected in examined soil. The most significant response of soil mycoflora was induced by the species identity of plant cover. Higher N fertilization significantly suppressed the abundance of soil microfungi at ambient CO₂. The effect of increased CO₂ on colony‐forming units was not significant when taken as an independent treatment; however, this factor interacted significantly with N availability. Some species, e.g. the Clonostachys rosea, were proven associated with the plant cover components, in this particular case with Trifolium repens. Therefore, we suggest the identity of plant species constituting plant cover as the most important factors affecting soil microfungi in agroecosystems.     

Nitrogen-regulated effects of free-air CO2 enrichment on methane emissions from paddy rice fields

Using the free‐air CO₂ enrichment (FACE) techniques, we carried out a 3‐year mono‐factorial experiment in temperate paddy rice fields of Japan (1998–2000) and a 3‐year multifactorial experiment in subtropical paddy rice fields in the Yangtze River delta in China (2001–2003), to investigate the methane (CH₄) emissions in response to an elevated atmospheric CO₂ concentration (200±40 mmol mol^?1 higher than that in the ambient atmosphere). No significant effect of the elevated CO₂ upon seasonal accumulative CH₄ emissions was observed in the first rice season, but significant stimulatory effects (CH₄ increase ranging from 38% to 188%, with a mean of 88%) were observed in the second and third rice seasons in the fields with or without organic matter addition. The stimulatory effects of the elevated CO₂ upon seasonal accumulative CH₄ emissions were negatively correlated with the addition rates of decomposable organic carbon (P<0.05), but positively with the rates of nitrogen fertilizers applied in either the current rice season (P<0.05) or the whole year (P<0.01). Six mechanisms were proposed to explain collectively the observations. Soil nitrogen availability was identified as an important regulator. The effect of soil nitrogen availability on the observed relation between elevated CO₂ and CH₄ emission can be explained by (a) modifying the C/N ratio of the plant residues formed in the previous growing season(s); (b) changing the inhibitory effect of high C/N ratio on plant residue decomposition in the current growing season; and (c) altering the stimulatory effects of CO₂ enrichment upon plant growth, as well as nitrogen uptake in the current growing season. This study implies that the concurrent enrichment of reactive nitrogen in the global ecosystems may accelerate the increase of atmospheric methane by initiating a stimulatory effect of the ongoing dramatic atmospheric CO₂ enrichment upon methane emissions from nitrogen‐poor paddy rice ecosystems and further amplifying the existing stimulatory effect in nitrogen‐rich paddy rice ecosystems.     

Carbon partitioning and rhizosphere C-flow in Lolium perenne as affected by CO2 concentration, irradiance and below-ground conditions

Plant responses to increasing atmospheric CO₂ concentrations have received considerable interest. However, major uncertainties in relation to interactive effects of CO₂ with above- and below-ground conditions remain. This microcosm study investigated the impacts of CO₂ concentration on plant growth, dry matter partitioning and rhizodeposition as affected by: (i) photon flux density (PFD), and (ii) growth matrix. Plants were grown in a sandy loam soil for 28 d under two photon flux densities: 350 (low PFD) and 1000 μmol m^–2 s^–1 (high PFD) and two CO₂ concentrations: 450 (low CO₂) and 720 μmol mol^–1 (high CO₂). Partitioning of recent assimilate amongst plant and rhizosphere C-pools was determined by use of ¹⁴CO₂ pulse-labelling. In treatments with high PFD and/or high CO₂, significant (P < 0.05) increases in dry matter production were found in comparison with the low PFD/low CO₂ treatment. In addition, significant (P < 0.05) reductions in shoot %N and SLA were found in treatments imposing high PFD and/or high CO₂. Root weight ratio (RWR) was unaffected by CO₂ concentration, however, partitioning of ¹⁴C to below ground pools was significantly (P < 0.05) increased. In a separate study, L. perenne was grown for 28 d in microcosms percolated with nutrient solution, in either a sterile sand matrix or nonsterile soil, under high or low CO₂. Dry matter production was significantly (P < 0.01) increased for both sand and soil grown seedlings. Dry matter partitioning was affected by matrix type. ¹⁴C-allocation below ground was increased for sand grown plants. Rhizodeposition was affected by CO₂ concentration for growth in each matrix, but was increased for plants grown in the soil matrix, and decreased for those in sand. The results illustrate that plant responses to CO₂ are potentially affected by (i) PFD, and (ii) by feedbacks from the growth matrix. Such feedbacks are discussed in relation to soil nutrient status and interactions with the rhizosphere microbial biomass.     

Biomass allocation in old-field annual species grown in elevated CO2 environments: no evidence for optimal partitioning

Increased atmospheric carbon dioxide supply is predicted to alter plant growth and biomass allocation patterns. It is not clear whether changes in biomass allocation reflect optimal partitioning or whether they are a direct effect of increased growth rates. Plasticity in growth and biomass allocation patterns was investigated at two concentrations of CO₂ ([CO₂]) and at limiting and nonlimiting nutrient levels for four fast‐ growing old‐field annual species. Abutilon theophrasti, Amaranthus retroflexus, Chenopodium album, and Polygonum pensylvanicum were grown from seed in controlled growth chamber conditions at current (350 μmol mol^?1, ambient) and future‐ predicted (700 μmol mol^?1, elevated) CO₂ levels. Frequent harvests were used to determine growth and biomass allocation responses of these plants throughout vegetative development. Under nonlimiting nutrient conditions, whole plant growth was increased greatly under elevated [CO₂] for three C3 species and moderately increased for a C4 species (Amaranthus). No significant increases in whole plant growth were observed under limiting nutrient conditions. Plants grown in elevated [CO₂] had lower or unchanged root:shoot ratios, contrary to what would be expected by optimal partitioning theory. These differences disappeared when allometric plots of the same data were analysed, indicating that CO₂‐induced differences in root:shoot allocation were a consequence of accelerated growth and development rates. Allocation to leaf area was unaffected by atmospheric [CO₂] for these species. The general lack of biomass allocation responses to [CO₂] availability is in stark contrast with known responses of these species to light and nutrient gradients. We conclude that biomass allocation responses to elevated atmospheric [CO₂] are not consistent with optimal partitioning predictions.