The influence of soil-type,drought and nitrogen addition on interactions between Calluna vulgaris and Deschampsia flexuosa: implications for heathland regeneration

Increasing dominance of grass species such as Deschampsia flexuosa on lowland dry heathlands is a major problem for land managers. This problem is especially acute in north-west Europe where replacement of dwarf-shrubs by grasses has been linked to high levels of nitrogen deposition. Interactions between Calluna vulgaris and D. flexuosa were examined using pot experiments under a variety of watering (drought vs. no drought), nitrogen addition (10 vs. 50 kg N ha^–1 yr^–1) and soil type (peaty vs. sandy) regimes. Two experiments using identical replacement series designs were used to investigate interactions between these two species at the seedling and young plant stage. Response of the two species to the environmental variables was different for seedlings and young plants. In the seedling experiment D. flexuosa responded significantly to soil type only, with greater cover on peaty soil than on sandy soil after 6 months. C. vulgaris germination showed a significant response to watering regime, with fewer seedlings in plots that had a drought treatment imposed. When the experiment was repeated using young plants of C. vulgaris and D. flexuosa both species responded significantly to soil type only. Height, % cover and shoot biomass of both species were greater when grown in peaty soil than when grown in sandy soil. Watering and nitrogen addition had few significant effects. Species interactions were mainly one-way. C. vulgaris biomass yield was reduced when grown in mixture with D. flexuosa, whilst D. flexuosa yield was unaffected by the presence of C. vulgaris. The yield density curve for D. flexuosa indicated that intra-specific competition was occurring at planting densities above 29 plants m^–2 whilst this was not seen for C. vulgaris. The effect of soil type on species interactions was also investigated. C. vulgaris was best able to compete with D. flexuosa on sandy soils where the shoot biomass ratio was greatest (C. vulgaris:D. flexuosa= 1:4.4). Differences in the responses of these species to the environmental variables investigated may help to explain heathland vegetation changes seen under field conditions.     

Effects of soil organic matter and nitrogen supply on competition between Festuca ovina and Deschampsia flexuosa during inland dune succession

We tested the prediction that the successional replacement of plant species during succession on inland sand dunes results from the effects of an increase in nitrogen mineralization on competitive interactions. The growth and competitive strength of Festuca ovina and Deschampsia flexuosa on soil substrates with different amounts of soil organic matter or nitrogen supply were measured. Small tillers of Festuca ovina and Deschampsia flexuosa were grown in monocultures and 1:1-mixtures on soil columns with undisturbed layers of soil organic matter from different successional age. There was (a) no visible soil organic matter, (b) a thin soil organic layer (0.5 cm) and (c) a thicker soil organic layer (6.0 cm) present on the soil columns. The species were also grown on columns with no visible soil organic matter (bare sand) with two different levels of N fertilization to mimic the increased N mineralization in the older successional stages.In monoculture, Festuca produced more biomass on the substrates with a soil organic layer compared to the unfertilized sand substrate. It also produced more biomass on sand substrates with N fertilization. Deschampsia produced more biomass in treatments with a soil organic layer compared to the bare sand treatments, but did not respond to the ammonium-nitrate addition. In competition, Festuca seemed to be the stronger competitor on the unfertilized sand substrate. Festuca was also the better competitor on the N fertilized sand treatments, while on the treatments with a soil organic layer Deschampsia was the winning species. Our results do not support the hypothesis that an increase in N supply is responsible for the replacement of Festuca by Deschampsia that concur with the accumulation of soil organic matter during succession in inland dunes.     

Effects of nitrogen addition on the stress sensitivity of Calluna vulgaris

The effects of low levels of nitrogen addition (7·7 and 15·4 kg N ha⁻¹ yr⁻¹) on plant sensitivity to biotic and abiotic stress were studied at a lowland heath in the south of England that has received N treatments since 1989. Larval growth rates and adult weights of heather beetles were found to be significantly higher when insects were reared on plants that had received additional N, with implications for insect survival and reproductive success. Electrolyte leakage measurements failed to reveal any significant impact of N addition on plant sensitivity to frost episodes in early winter. In April, however, there was some evidence of slightly decreased frost hardiness in plants receiving additional N. Accelerated spring bud burst also suggested earlier physiological activity in N-treated plots. The rate of water loss from excised shoots of Calluna vulgaris (L.) Hull was significantly faster in plants receiving additional N, although no difference in plant water potential was measured in the field after a prolonged dry spell. Whilst experimental N addition had only a small effect on plant sensitivity to abiotic stress, the relationship between enhanced deposition and increased insect performance was clear, with the potential for substantially increased insect damage at deposition rates around the critical load contributing to the formation of gaps in the Calluna canopy.     

Effects of increased temperature and nitrogen availability on the displacement of Calluna vulgaris by Pteridium aquilinum

Abstract. Atmospheric greenhouse gas concentrations will continue to rise for the foreseeable future, and, due to radiative forcing, surface temperatures will also increase. If soils warm sufficiently, this will result in increased litter decomposition rates, while nitrogen availability will also rise due to increased deposition. A field-based experiment manipulating air temperatures using polythene tents and nitrogen availability by adding 50 kg N ha^-1 yr^-1 has shown that this will result in further encroachment of Pteridium into Calluna moorland. Although this experiment, treating plots on and around the boundary between Pteridium and Calluna, did not show any statistically significant effect on the movement of the boundary, there were clear indications of Calluna dieback at the boundary. In addition, the frequency of Pteridium fronds increased in the Calluna-dominated areas, while Calluna became less abundant in the pure Pteridium plots. Calluna dieback was not due to litter accumulation as is frequently assumed, but the dieback is likely to be caused by shading of Calluna by the increased growth of Pteridium fronds. Botanical diversity was higher in the Pteridium plots, but considering the importance of grouse shooting for the maintenance of Calluna moorland, displacement of Calluna by Pteridium may result in a lower conservation and economic value of moorlands.     

Leaf growth in the fast-growing Holcus lanatus and the slow-growing Deschampsia flexuosa: tissue maturation

Inherent variation in the relative growth rate of grasses is negatively correlated with that in leaf mass per unit leaf area (LMA). To scrutinize this correlation, the LMA of two grass species was analysed. Changes in LMA and cell wall synthesis in leaf blades of the fast-growing grass Holcus lanatus and the slow-growing grass Deschampsia flexuosa were investigated above the elongation zone of the leaf blade. After the leaf had obtained its final length, in H. lantus final LMA values of 40-44 gm^-2 were obtained at full leaf length, whereas in D. flexuosa LMA values continued to rise to 110 gm^-2. During this period of tissue maturation the LMA value doubled in H. lanatus, whereas in D. flexuosa an increase of 30% was measured. Most of the cell walls could be hydrolysed with driselase, the residue was hydrolysed with sulphuric acid. Driselase hydrolysates were identical in sugar composition, whereas the sugars released by sulphuric acid treatment changed gradually in composition as the tissue matured. The major sites of cell wall deposition during cell maturation were the outer walls of epidermal cells, fibres adjacent to the epidermis and the mestome ring around the vascular bundles. Lignin deposition was restricted to the vascular bundles and lignin levels of the leaf blade did not exceed 0.9% of the total amount of cell wall polysaccharides. Lignin accumulation occurred mainly after the increase in LMA and is unlikely to affect measurably the growth of these leaves.     

The role of different arbuscular mycorrhizal fungi in the growth of Calamagrostis villosa and Deschampsia flexuosa, in experiments with simulated acid rain

A series of microcosm experiments was established to investigate the effects of simulated acid rain on the capacity of three arbuscular mycorrhizal fungi (AMF) to germinate and colonize two grasses, Calamagrostis villosa and Deschampsia flexuosa. These two grasses are normally found in degraded Norway spruce forests in the Northern Czech Republic where acid rain pollution exists and C. villosa initially outcompetes D. flexuosa for the same niche. An AM fungus isolated from acid soils (Acaulospora tuberculata BEG41) was more tolerant of acidification than two species of Glomus (isolated from agricultural soils of neutral pH) in microcosm studies. Different effects of simulated acid rain (SAR) were found at all stages of the development of three AMF studied in model systems, including spore germination, colonization of host roots, and alkaline phosphatase (ALP) and NADH diaphorase activity of the extraradical mycelium. No ALP activity was found in hyphae germinating from the spores without plants whereas it was detected in all hyphae linked to a functioning intraradical mycelium.Simulated acid rain also affected the mycorrhizal growth response and belowground competition of the two grasses. Disturbance of the ERM between the two plant species significantly reduced the growth of C. villosa but not D. flexuosa. Disturbance also decreased root colonization by AMF of both plants, the total length of ERM and the total length of extraradical hyphae with ALP and NADH diaphorase activity adjacent to both plants. D. flexuosa appeared less dependent on the mycorrhizal state, for shoot and root growth, than C. villosa under the experimental conditions. The ability, therefore, of C. villosa to thrive in forest stands suffering from acid rain pollution may be related to this dependence on its mycorrhizal hyphal links to D. flexuosa under the environmental conditions produced by the pollution including higher light levels.     

彩萼石楠的组织培养和快速繁殖

1植物名称彩萼石楠或帚石南(Calluna vulgaris). 2材料类别当年生新梢. 3培养条件基本培养基为MS.(1)诱导芽培养基:MS 6-BA 0.5～2.0 mg·L-1(单位下同) NAA 0.05;(2)增殖培养基:MS 6-BA 1.0～2.0 IAA 0.2 GA 0.5;(3)生根培养基:1/2MS IBA 0.5～1.0.以上培养基均附加30 g·L-1蔗糖、6 g·L-1琼脂,pH 5.4～5.8.光照度为1 000～2 500 lx,光照时间12 h·d-1;培养温度为(25±1)℃.     

Partitioning of soil phosphorus regulates competition between Vaccinium vitis‐idaea and Deschampsia cespitosa

It has been hypothesized that the wide range of forms and complexities of phosphorus (P) in soil may result in resource partitioning that contributes to the maintenance of plant species diversity. Here, we test whether the graminoid, Deschampsia cespitosa, and the ericaceous shrub, Vaccinium vitis‐idaea, which often coexist, display preferences in utilization of P forms, and differ in their production of extracellular P‐degrading enzymes. We provided plants with no additional P, or P forms with decreasing lability, namely sodium phosphate (SP), D‐glucose 6 phosphate (DG6P), sodium phytate (PASS), and a combination of SP, DG6P, and PASS. We also tested if preferences for P forms affected the competitive outcomes between the two species compared between conspecifics, as indicated by shoot biomass and acquisition of nitrogen (N) and P. Both D. cespitosa and V. vitis‐idaea produced the greatest biomass when supplied with a mix of all three forms of P. Of the three forms of P tested alone, shoot biomass produced by both species was least when supplied with SP. D. cespitosa performed better when grown with PASS or a mix of all P forms compared with the performance of V. vitis‐idaea on these substrates. This was reflected by substantially greater phytase activity on the surface of its roots compared with V. vitis‐idaea. In contrast, V. vitis‐idaea produced more phosphomonoesterase to hydrolyze the simple organic P form, DG6P. Although N was kept constant in the treatments, the ability of plants to acquire it was dependent on species identity, competition, and P supply. These findings provide direct evidence for preferences toward specific forms of P and indicate a key role played by organic forms of P. The results support the idea that partitioning for soil P is one factor regulating plant competition, and ultimately, community composition. Our data also highlight the importance of the interplay between P supply and N acquisition.     

Influence of arbuscular mycorrhizal fungi and simulated acid rain on the growth and coexistence of the grasses Calamagrostis villosa and Deschampsia flexuosa

Two grass species — Calamagrostis villosa (Chaix) J.F. Gmelin and Deschampsia flexuosa (L.) Trin. — are expanding in mountain Norway spruce (Picea abies L. Karst.) forests of Central Europe damaged by anthropogenic pollution constituted particularly of acid rain. This invasion of grasses may be caused by the higher irradiance reaching the forest floor after the pollution-induced tree defoliation. The relative abundance of the two grass species is changing during the process of forest decline. Our study investigated the effects of arbuscular mycorrhizal fungi (AMF) on the growth and coexistence of both species under simulated acid rain (SAR) and two levels of irradiance. Three microcosm experiments were conducted to investigate how both grasses are influenced by the AMF when grown separately or together interacting via extraradical mycelium (ERM). A positive growth response to inoculation with Glomus mosseae BEG 25 was found for both grass species when cultivated separately and the mycorrhizal dependence and the growth benefit for D. flexuosa was greater than for C. villosa. However, when both grass species were grown together in the rhizoboxes with separated root and hyphal compartments, the growth effect of the AMF was the opposite, i.e. C. villosa benefited more. The plants did not benefit from the AMF inoculation under the SAR treatment compared with dH2O treatment. The SAR also negatively influenced root length colonised by AMF, length of the ERM, alkaline phosphatase and NADH diaphorase activities of the ERM. The role of the ERM in transporting phosphorus between these grasses was verified by applying the radioisotope ³²P. There was a greater transport of isotopic ³²P between inoculated plants C. villosa and D. flexuosa grown in separated root compartments, as compared to non-inoculated plants. The amount of transported ³²P was low: a maximum of 3% of applied ³²P was detected in the shoots of receiver plants. Mechanical disturbance of the ERM significantly decreased the ³² P transport between plants. The ³²P transport between mycorrhizal plants was higher in the D. flexuosa to C. villosa direction than in the opposite one. Neither the SAR nor the low level of irradiance influenced the amount of transported ³²P. We discuss the role of ERM links between root systems in the coexistence of both grass species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Latent soil effects of grazing and ammonium deposition on Deschampsia flexuosa tillers inserted and grown in heather moorland soil

Upland heather moorlands support a range of increasingly rare and threatened biota, making them a priority habitat for conservation and restoration. Over-grazing and nitrogen deposition are two of the most important threats to maintaining these heather moorlands, yet there remains a paucity of research into their combined long-term effects. During the summer of 2008, we established an experiment within an existing research site at Pwllpeiran, mid-Wales. The original site, established in 1996, investigated long-term grazing and N deposition treatments (ammonium and nitrate) on heather moorland. Previous findings from the site, following a decade of treatment, suggested no significant, detectable impacts of treatments on soil C:N ratio and the foliar nitrogen of vegetation.The aim of our study was to investigate short- (2008) and long-term (1996–2008) N deposition treatment impacts, in combination with past grazing (1990–2007), on soil nutrient bioavailability. Soil cores were harvested and aboveground vegetation removed. Tillers of the grass Deschampsia flexuosa were planted into these cores which were then reciprocally transplanted back into the field experiment. The N deposition treatment was continued but grazing was excluded. D. flexuosa biomass changes were next assessed and leaf chemistry investigated using the metabolic fingerprinting method Fourier-transform infrared spectroscopy (FTIR) following three months of growth in the field (May–August 2008).Grazing treatment (on its own) had significant negative impacts on aboveground biomass and significant changes in plant chemistry were also revealed through the metabolic fingerprinting method Fourier transform infrared spectroscopy (FTIR). Short-term N deposition treatments during 2008 had no impacts on D. flexuosa growth or chemistry. There were also no detectable latent effects of long-term nitrate treatments on either growth or chemistry of D. flexuosa. However, plants grown in plots that had received long-term treatments of ammonium (NH₄⁺) had significantly lower poly-phenolic contents (revealed by FTIR) than plants grown in either nitrate (NO₃⁻) or control plots, suggesting detectable latent effects of N application in its reduced form. Further work needs to be undertaken to assess the relevance of residual soil nitrogen pools post N deposition and grazing.     

Effects of salinity and temperature on Deschampsia antarctica

Deschampsia antarctica is one of two species of vascular plants native to Antarctica. Populations of D. antarctica have become established on recently exposed glacial forelands on the Antarctic Peninsula and these plants may rely upon nutrient inputs from hauled out mammals, seabirds and sea spray. However, not much is known about the ability of these plants to tolerate salinity stress. We examined the effects of salinity and temperature on growth, reproduction, chlorophyll fluorescence and water relations of D. antarctica. In addition, we analysed concentrations of free proline in leaves and roots as previous studies have found large increases in the concentration of this amino acid in response to environmental stress. The growth chamber experiment was a 3 × 3 (temperature × salinity) complete factorial. Plants were grown at three temperature regimes: 7°/7°C, 12°/7°C, and 20°/7°C day/night and three salinity levels: <0.02 decismen per metre (dS m⁻¹; “low salinity”), 2.5 dS m⁻¹ (“medium salinity”), and 5.0 dS m⁻¹ (“high salinity”) for 66 days. Warmer temperatures improved leaf and tiller production as well as leaf and root length, which is consistent with previous findings on this species. Salinity reduced final root length by 6 and 13% in the medium and high-salinity treatments, respectively. Plants growing in medium and high-salinity treatments had xylem pressures that were more negative and higher free-proline concentrations, suggesting that proline may act as an osmoregulant in D. antarctica.     

The Lepidoptera fauna associated with Calluna vulgaris: effects of plant architecture on abundance and diversity

Abstract .1. The effects of plant architecture on the Lepidoptera assemblage associated with heather Calluna vulgaris were studied at four locations in northern England and southern Scotland in 1992 and 1993. The study areas were Calluna vulgaris – Vaccinium myrtillus heaths, where management by rotational burning had created a mosaic of stands of different-aged Calluna . Lepidoptera were sampled in the larval stage.
2. Larval abundance was considered in relation to Calluna age, height, cover, green shoot density, and flower density. For most study areas and species, multiple regression selected Calluna height as the variable that explained most variation in larval density, although the explanatory variables were intercorrelated.
3. After compensating for the effects of different study locations, the slope of the regression between the logarithm of larval density and the logarithm of Calluna height was common to the macrolepidoptera, microlepidoptera, geometrid, and noctuid larval groups.
4. There was a significant progressive increase in larval diversity with increase in Calluna height, due to the presence of uncommon moth species in the samples from taller Calluna, and a change in the contribution of common species to the community in different height zones.
5. The significance of these results is discussed in relation to other heathland invertebrate studies and management practices.     

不同pH的酚酸溶液对杉木及萝卜幼苗生长的影响

通过四因素五水平的二次回归正交旋转试验设计 ,研究 pH值、肉桂酸、对羟基苯甲酸、香草醛四个因素对杉木和萝卜幼苗生长的影响。结果表明 ,杉木幼苗的鲜重生长随酚酸溶液的pH值的增大而增加 ,而随肉桂酸、对羟基苯甲酸和香草醛浓度的增大而减小。比较 3种不同酚类物质对杉木幼苗生长的影响 ,肉桂酸对杉木幼苗生长的影响 >对羟基苯甲酸 >香草醛 ,且 pH值与肉桂酸之间 ;肉桂酸与香草醛之间 ;对羟基苯甲酸与香草醛之间的交互作用都达到显著水平。萝卜幼苗生长对不同pH的酚酸溶液的反应与杉木幼苗相类似 ,随酚酸溶液的pH值的增大而增加 ,而随肉桂酸、对羟基苯甲酸和香草醛浓度的增大而减小。但对羟基苯甲酸对萝卜幼苗生长的影响 >肉桂酸 >香草醛 ,酚酸溶液的 pH值与所有 3种酚酸之间的交互作用都达到显著水平 ;肉桂酸与香草醛的交互作用对萝卜幼苗生长的影响也达到F0 0 1水平显著。     

Fungal associations of Danish Calluna vulgaris roots with special reference to ericoid mycorrhiza

Fungi were isolated from young, serial-washed roots of Calluna sampled from a Danish heathland, Hjelm Hede. Of the 626 isolates, those that were dark, sterile and septate were divided into 13 morphological groups based on their appearance in culture on malt agar. Mycorrhizal synthesis in vitro showed that several groups formed typical ericoid mycorrhiza with seedlings of Calluna; these ericoid mycorrhizal fungi were morphologically similar to Hymenoscyphus ericae. The identities of the other dark, septate fungi are uncertain. Oidiodendron spp. were isolated in a very low frequency; these fungi also formed typical ericoid mycorrhiza. The Calluna root system on Hjelm Hede demonstrated a high morphological diversity among the associated dark, septate fungi suggesting that more than one fungus could coexist in the same host root system.     

Influence of Nitrogen and Winter Climate Stresses on Calluna vulgaris(L.) Hull

Four nitrogen (N) levels, spanning those found naturally, wereestablished in Calluna vulgaris grown in containers in an oceanicclimate for 4 years on the southwest coast of Norway. EstablishedN contents of the plants were 0.7, 0.8, 1.3 and 2.5% of dryweight. Experiments comprising 5000 plants were performed inthe open and in growth chambers. In the open, dry matter accumulationand date of bud burst were recorded at the different N levels.Breaking of winter dormancy was studied under controlled conditions.In October/November, plants needed 1474 day-degrees for budburst at the lowest N level, whereas at the highest level only582 day-degrees were required. The natural start of growth inthe field was on 14 April at the highest N level and on 26 Aprilat the lowest. Freeze testing of the shoots (with the rootsprotected from frost) revealed that Calluna tolerates temperaturesas low as -15 °C as late as the beginning of April. However,plants at the two medium N levels were most tolerant of frost.During the last winter of the experiments, nearly 50% of plantsin the open died at the highest N level, whereas plants at thelower levels survived without damage. The effects of N are discussedin relation to the influence on physiological ageing of theplants, the change in nutrient balance of the plant tissue andthe subsequent increase in susceptibility to biotic and abioticstresses. Copyright 2001 Annals of Botany Company Calluna vulgaris, climate stress, growth initiation, nitrogen     

Growth pattern and distribution of biomass of Calluna vulgaris on an ombrotrophic peat bog

The distribution pattern of Calluna vulgaris on the ombrotrophic peat bog Åkhultmyren, S. Sweden, was investigated.
Calluna occurs primarily on hummocks, with its maximum biomass (ca 250 g m⁻² above ground, and ca 300 g m⁻² below ground) concentrated on hummock tops. Biomass decreased gradually along the hummock-hollow gradient to a level of ca 5 cm above the water level. This decreasing biomass was correlated with a decrease in the predominant age of the above ground shoots from 6–8 yr to 3–4 yr old shoots.
Calluna is characterized by extremely slow diameter growth and biomass increment. There was, however, a poor relation between height above water level and stem thickness, assimilating biomass and total shoot weight, respectively.     

Competition for nutrients between Calluna vulgaris (L.) Hull and Molinia caerulea (L.) Moench

Summary A combination of a removal and fertilization experiment in the field and a replacement experiment in containers in a cold frame was carried out to investigate the role of nutrient supply in the interactions between Calluna vulgaris and Molinia caerulea. It is concluded that the growth of Calluna as well as that of Molinia increased with increasing nutrient availability. However, the increase in biomass of Molinia was much larger than that of Calluna. It is also concluded that increased nutrient availability favours the competitive vigour of Molinia in interaction with Calluna.     

Interactive effects of elevated CO2, warming, and drought on photosynthesis of Deschampsia flexuosa in a temperate heath ecosystem

Global change factors affect plant carbon uptake in concert. In order to investigate the response directions and potential interactive effects, and to understand the underlying mechanisms, multifactor experiments are needed. The focus of this study was on the photosynthetic response to elevated CO(2) [CO2; free air CO(2) enrichment (FACE)], drought (D; water-excluding curtains), and night-time warming (T; infrared-reflective curtains) in a temperate heath. A/C(i) curves were measured, allowing analysis of light-saturated net photosynthesis (P(n)), light- and CO(2)-saturated net photosynthesis (P(max)), stomatal conductance (g(s)), the maximal rate of Rubisco carboxylation (V(cmax)), and the maximal rate of ribulose bisphosphate (RuBP) regeneration (J(max)) along with leaf δ(13)C, and carbon and nitrogen concentration on a monthly basis in the grass Deschampsia flexuosa. Seasonal drought reduced P(n) via g(s), but severe (experimental) drought decreased P(n) via a reduction in photosynthetic capacity (P(max), J(max), and V(cmax)). The effects were completely reversed by rewetting and stimulated P(n) via photosynthetic capacity stimulation. Warming increased early and late season P(n) via higher P(max) and J(max). Elevated CO(2) did not decrease g(s), but stimulated P(n) via increased C(i). The T×CO2 synergistically increased plant carbon uptake via photosynthetic capacity up-regulation in early season and by better access to water after rewetting. The effects of the combination of drought and elevated CO(2) depended on soil water availability, with additive effects when the soil water content was low and D×CO2 synergistic stimulation of P(n) after rewetting. The photosynthetic responses appeared to be highly influenced by growth pattern. The grass has opportunistic water consumption, and a biphasic growth pattern allowing for leaf dieback at low soil water availability followed by rapid re-growth of active leaves when rewetted and possibly a large resource allocation capability mediated by the rhizome. This growth characteristic allowed for the photosynthetic capacity up-regulations that mediated the T×CO2 and D×CO2 synergistic effects on photosynthesis. These are clearly advantageous characteristics when exposed to climate changes. In conclusion, after 1 year of experimentation, the limitations by low soil water availability and stimulation in early and late season by warming clearly structure and interact with the photosynthetic response to elevated CO(2) in this grassland species.