Influence of elevated CO2 and nitrogen nutrition on rice plant growth,soil microbial biomass,dissolved organic carbon and dissolved CH4

Rice (Oryza sativa) was grown in six sunlit, semi-closed growth chambers for two seasons at 350 L L^–1 (ambient) and 650 L L^–1 (elevated) CO₂ and different levels of nitrogen (N) supplement. The objective of this research was to study the influence of CO₂ enrichment and N nutrition on rice plant growth, soil microbial biomass, dissolved organic carbon (DOC) and dissolved CH₄. Elevated CO₂ concentration ([CO₂]) demonstrated a wide range of enhancement to both above- and below-ground plant biomass, in particular to stems and roots (for roots when N was not limiting) in the mid-season (80 days after transplanting) and stems/ears at the final harvest, depending on season and the level of N supplement. Elevated [CO₂] significantly increased microbial biomass carbon in the surface 5 cm soil when N (90 kg ha^–1) was in sufficient supply. Low N supplement (30 kg ha^–1) limited the enhancement of root growth by elevated [CO₂], leading consequently to diminished response of soil microbial biomass carbon to CO₂ enrichment. The concentration of dissolved CH₄ (as well as soil DOC, but to a lesser degree) was observed to be positively related to elevated [CO₂], especially at high rate of N application (120 kg ha^–1) or at 10 cm depth (versus 5 cm depth) in the later half of the growing season (at 80 kg N ha^–1). Root senescence in the late season complicated the assessment of the effect of elevated [CO₂] on root growth and soil organic carbon turnover and thus caution should be taken when interpreting respective high CO₂ results.     

Plant Nitrogen Dynamics in Shortgrass Steppe under Elevated Atmospheric Carbon Dioxide

The direct and indirect effects of increasing levels of atmospheric carbon dioxide (CO₂) on plant nitrogen (N) content were studied in a shortgrass steppe ecosystem in northeastern Colorado, USA. Beginning in 1997 nine experimental plots were established: three open-top chambers with ambient CO₂ levels (approximately 365 mol mol^–1), three open-top chambers with twice-ambient CO₂ levels (approximately 720 mol mol^–1), and three unchambered control plots. After 3 years of growing-season CO₂ treatment, the aboveground N concentration of plants grown under elevated atmospheric CO₂ decreased, and the carbon–nitrogen (C:N) ratio increased. At the same time, increased aboveground biomass production under elevated atmospheric CO₂ conditions increased the net transfer of N out of the soil of elevated-CO₂ plots. Aboveground biomass production after simulated herbivory was also greater under elevated CO₂ compared to ambient CO₂. Surprisingly, no significant changes in belowground plant tissue N content were detected in response to elevated CO₂. Measurements of individual species at peak standing phytomass showed significant effects of CO₂ treatment on aboveground plant tissue N concentration and significant differences between species in N concentration, suggesting that changes in species composition under elevated CO₂ will contribute to overall changes in nutrient cycling. Changes in plant N content, driven by changes in aboveground plant N concentration, could have important consequences for biogeochemical cycling rates and the long-term productivity of the shortgrass steppe as atmospheric CO₂ concentrations increase.     

Interactive effects of soil nitrogen and atmospheric carbon dioxide on root/rhizosphere carbon dioxide efflux from loblolly and ponderosa pine seedlings

We measured CO₂ efflux from intact root/rhizosphere systems of 155 day old loblolly (Pinus taeda L.) and ponderosa (Pinus ponderosa Dougl. ex Laws.) pine seedlings in order to study the effects of elevated atmospheric CO₂ on the below-ground carbon balance of coniferous tree seedlings. Seedlings were grown in sterilized sand culture, watered daily with either 1, 3.5 or 7 mt M NH ₄ ⁺ , and maintained in an atmosphere of either 35 or 70 Pa CO₂. Carbon dioxide efflux (mol CO₂ plant^–1 s^–1) from the root/rhizosphere system of both species significantly increased when seedlings were grown in elevated CO₂, primarily due to large increases in root mass. Specific CO₂ efflux (mol CO₂ g root^–1 s^–1) responded to CO₂ only under conditions of adequate soil nitrogen availability (3.5 mt M). Under these conditions, CO₂ efflux rates from loblolly pine increased 70% from 0.0089 to 0.0151 mol g^–1 s^–1 with elevated CO₂ while ponderosa pine responded with a 59% decrease, from 0.0187 to 0.0077 mol g^–1 s^–1. Although below ground CO₂ efflux from seedlings grown in either sub-optimal (1 mt M) or supra-optimal (7 mt M) nitrogen availability did not respond to CO₂, there was a significant nitrogen treatment effect. Seedlings grown in supra-optimal soil nitrogen had significantly increased specific CO₂ efflux rates, and significantly lower total biomass compared to either of the other two nitrogen treatments. These results indicate that carbon losses from the root/rhizosphere systems are responsive to environmental resource availability, that the magnitude and direction of these responses are species dependent, and may lead to significantly different effects on whole plant carbon balance of these two forest tree species.     

Elevated CO2 effects on carbon and nitrogen cycling in grass/clover turves of a Psammaquent soil

Effects of elevated CO₂ (525 and 700 L L^–1), and a control (350 L L^–1 CO₂), on biochemical properties of a Mollic Psammaquent soil in a well-established pasture of C3 and C4 grasses and clover were investigated with continuously moist turves in growth chambers over four consecutive seasonal temperature regimes from spring to winter inclusive. After a further spring period, half of the turves under 350 and 700 L L^–1 were subjected to summer drying and were then re-wetted before a further autumn period; the remaining turves were kept continuously moist throughout these additional three consecutive seasons. The continuously moist turves were then pulse-labelled with ¹⁴C-CO₂ to follow C pathways in the plant/soil system during 35 days.Growth rates of herbage during the first four seasons averaged 4.6 g m^–2 day^–1 under 700 L L^–1 CO₂ and were about 10% higher than under the other two treatments. Below-ground net productivity at the end of these seasons averaged 465, 800 and 824 g m^–2 in the control, 525 and 700 L L^–1 treatments, respectively.in continuously moist soil, elevated CO₂ had no overall effects on total, extractable or microbial C and N, or invertase activity, but resulted in increased CO₂-C production from soil, and from added herbage during the initial stages of decomposition over 21 days; rates of root decomposition were unaffected. CO₂ produced h^–1 mg^–1 microbial C was about 10% higher in the 700 L L^–1 CO₂ treatment than in the other two treatments. Elevated CO₂ had no clearly defined effects on N availability, or on the net N mineralization of added herbage.In the labelling experiment, relatively more ¹⁴C in the plant/soil system occurred below ground under elevated CO₂, with enhanced turnover of ¹⁴C also being suggested.Drying increased levels of extractable C and organic-N, but decreased mineral-N concentrations; it had no effect on microbial C, but resulted in lowered microbial N in the control only. In soil that had been previously summer-dried, CO₂ production was again higher, but net N mineralization was lower, under elevated CO₂ than in the control after autumn pasture growth.Over the trial period of 422 days, elevated CO₂ generally appears to have had a greater effect on soil C turnover than on soil C pools in this pasture ecosystem.     

Elevated CO2 and temperature effects on soil carbon and nitrogen cycling in ryegrass/white clover turves of an Endoaquept soil

Effects of elevated CO₂ (700 L L^–1) and a control (350 L L^–1 CO₂) on the productivity of a 3-year-old ryegrass/white clover pasture, and on soil biochemical properties, were investigated with turves of a Typic Endoaquept soil in growth chambers. Temperature treatments corresponding to average winter, spring, and summer conditions in the field were applied consecutively to all of the turves. An additional treatment, at 700 L L^–1 CO₂ and a temperature 6°C higher throughout than in the other treatments, was included.Under the same temperature conditions, overall herbage yields in the 700 L L^–1 CO₂ treatment were ca. 7% greater than in the control at the end of the summer period. Root mass (to ca 25 cm depth) in the 700 L L^–1 CO₂ treatment was then about 50% greater than in the control, but in the 700 L L^–1 CO₂+6°C treatment it was 6% lower than in the control. Based on decomposition results, herbage from the 700 L L^–1+6°C treatment probably contained the highest proportion of readily decomposable components.Elevated CO₂ had no consistent effect on soil total C and N, microbial C and N, or extractable C concentrations in any of the treatments. Under the same temperature conditions, it did, however, enhance soil respiration (CO₂-C production) and invertase activity. The effects of elevated CO₂ on rates of net N mineralization were less distinct, and the apparent availability of N for the sward was not affected. Under elevated CO₂, soil in the higher-temperature treatment had a higher microbial C:N ratio; it also had a greater potential to degrade plant materials.Data interpretation was complicated by soil spatial variability and the moderately high background levels of organic matter and biochemical properties that are typical of New Zealand pasture soils. More rapid cycling of C under CO₂ enrichment is, nevertheless, indicated. Futher long-term experiments are required to determine the overall effect of elevated CO₂ on the soil C balance.     

Soil respiration response to three years of elevated CO2 and N fertilization in ponderosa pine (Pinus ponderosa Doug. ex Laws.)

We measured growing season soil CO₂ evolution under elevated atmospheric [CO₂] and soil nitrogen (N) additions. Our objectives were to determine treatment effects, quantify seasonal variation, and compare two measurement techniques. Elevated [CO₂] treatments were applied in open-top chambers containing ponderosa pine (Pinus ponderosa L.) seedlings. N applications were made annually in early spring. The experimental design was a replicated factorial combination of CO₂ (ambient, + 175, and +350 L L^–1 CO₂) and N (0, 10, and 20 g m^–2 N as ammonium sulphate). Soils were irrigated to maintain soil moisture at > 25 percent. Soil CO₂ evolution was measured over diurnal periods (20–22 hours) in October 1992, and April, June, and October 1993 and 1994 using a flow-through, infrared gas analyzer measurement system and corresponding pCO₂ measurements were made with gas wells. Significantly higher soil CO₂ evolution was observed in the elevated CO₂ treatments; N effects were not significant. Averaged across all measurement periods, fluxes, were 4.8, 8.0, and 6.5 for ambient + 175 CO₂, and +350 CO₂ respectively).Treatment variation was linearly related to fungal occurrence as observed in minirhizotron tubes. Seasonal variation in soil CO₂ evolution was non-linearly related to soil temperature; i.e., fluxes increased up to approximately soil temperature (10cm soil depth) and decreased dramatically at temperatures > 18°C. These patterns indicate exceeding optimal temperatures for biological activity. The dynamic, flow-through measurement system was weakly correlated (r = 0.57; p < 0.0001; n = 56) with the pCO₂ measurement method.     

Leaf cavity CO2 concentrations and CO2 exchange in onion,Allium cepa L.

Onion (Allium cepa L.) plants were examined to determine the photosynthetic role of CO₂ that accumulates within their leaf cavities. Leaf cavity CO₂ concentrations ranged from 2250 L L^–1 near the leaf base to below atmospheric (<350 L L^–1) near the leaf tip at midday. There was a daily fluctuation in the leaf cavity CO₂ concentrations with minimum values near midday and maximum values at night. Conductance to CO₂ from the leaf cavity ranged from 24 to 202 mol m^–2 s^–1 and was even lower for membranes of bulb scales. The capacity for onion leaves to recycle leaf cavity CO₂ was poor, only 0.2 to 2.2% of leaf photosynthesis based either on measured CO₂ concentrations and conductance values or as measured directly by ¹⁴CO₂ labeling experiments. The photosynthetic responses to CO₂ and O₂ were measured to determine whether onion leaves exhibited a typical C₃-type response. A linear increase in CO₂ uptake was observed in intact leaves up to 315 L L^–1 of external CO₂ and, at this external CO₂ concentration, uptake was inhibited 35.4±0.9% by 210 mL L^–1 O₂ compared to 20 mL L^–1 O₂. Scanning electron micrographs of the leaf cavity wall revealed degenerated tissue covered by a membrane. Onion leaf cavity membranes apparently are highly impermeable to CO₂ and greatly restrict the refixation of leaf cavity CO₂ by photosynthetic tissue.Abbreviations C_a external CO₂ concentration - C_i intercellular CO₂ concentration - CO₂ compensation concentration - PPFR photosynthetic photon fluence rate     

Long-term effects of CO2 enrichment and temperature increase on a temperate grass sward

Perennial ryegrass swards were grown in large containers on a soil and were exposed during two years to elevated (700 L L^-1) or ambient atmospheric CO₂ concentration at outdoor temperature and to a 3 °C increase in air temperature in elevated CO₂. The nitrogen nutrition of the grass sward was studied at two sub-optimal (160 and 530 kg N ha^-1 y^-1) and one non-limiting (1000 kg N ha^-1 y^-1) N fertilizer supplies. At cutting date, elevated CO₂ reduced by 25 to 33%, on average, the leaf N concentration per unit mass. Due to an increase in the leaf blade weight per unit area in elevated CO₂, this decline did not translate for all cuts in a lower N concentration per unit leaf blade area. With the non-limiting N fertilizer supply, the leaf N concentration (% N) declined with the shoot dry-matter (DM) according to highly significant power models in ambient (% N=4.9 DM^-0.38) and in elevated (%N=5.3 DM^-0.52) CO₂. The difference between both regressions was significant and indicated a lower critical leaf N concentration in elevated than in ambient CO₂ for high, but not for low values of shoot biomass. With the sub-optimal N fertilizer supplies, the nitrogen nutrition index of the grass sward, calculated as the ratio of the actual to the critical leaf N concentration, was significantly lowered in elevated CO₂. This indicated a lower inorganic N availability for the grass plants in elevated CO₂, which was also apparent from the significant declines in the annual nitrogen yield of the grass sward and in the nitrate leaching during winter. For most cuts, the harvested fraction of the plant dry-matter decreased in elevated CO₂ due, on average, to a 45–52% increase in the root phytomass. In the same way, a smaller share of the plant total nitrogen was harvested by cutting, due, on average, to a 25–41% increase in the N content of roots. The annual means of the DM and N harvest indices were highly correlated to the annual means of the nitrogen nutrition index. Changes in the harvest index and in the nitrogen nutrition index between ambient and elevated CO₂ were also positively correlated. The possible implication of changes in the soil introgen cycle and of a limitation in the shoot growth potential of the grass in elevated. CO₂ is discussed.Abbreviations 350 outdoor climate - 700 outdoor climate +350 L L^-1[CO₂] - 700+ outdoor climate +350 L L^-1 (CO₂) and +3 °C - N-- low N fertilizer supply - N+ high N fertilizer supply - N++ non-limiting N fertilizer supply - DM dry-matter     

Effects of <Emphasis Type="Italic">in vitro</Emphasis>rooting environments and irradiance on growth and photosynthesis of strawberry plantlets during acclimatization

Strawberry (Fragaria ananassaDuch. cv. Fengxiang) plantlets were cultured under two in vitroenvironments for rooting, and then acclimatized under two ex vitroirradiance conditions. At the end of rooting stage plant height, fresh weight and specific leaf area of T1-plants grown under high sucrose concentration (3 sucrose), low photosynthetic photon flux density (30 mol m^–2 s^–1) and normal CO₂ concentration (350–400 l l^–1) were significantly higher than those of T2-plantlets grown under low sucrose concentration (0.5), high photosynthetic photon flux density (90 mol m^–2 s^–1) and elevated CO₂ concentration (700–800 l l^–1). But T2-plantlets had higher net photosynthetic rate (Pn), effective photochemical quantum yield of PSII (_PSII), effective photosynthetic electron transport rate (ETR), photochemical quenching (q_P) and ratio of chlorophyll fluorescence yield decrease (Rfd). After transfer, higher irradiance obviously promoted the growth of plantlets and was beneficial for the development of photosynthetic functions during acclimatization. T2-plantlets had higher fresh weight, leaf area, _PSII and ETR under higher ex vitroirradiance condition.     

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.     

Effects of atmospheric CO2 enrichment, water status and applied nitrogen on water- and nitrogen-use efficiencies of wheat

Atmospheric CO₂ levels are expected to exceed 700 mol mol^–1 by the end of the 21st century. The influence of increased CO₂ concentration on crop plants is of major concern. This study investigated water- and nitrogen-use efficiency (WUE and NUE, respectively, were defined by the amount of biomass accumulated per unit water or N uptake) of spring wheat (Triticum aestivumL.) grown under two atmospheric CO₂ concentrations (350 and 700 mol mol^–1), two soil moisture treatments (well-watered and drought) and five nitrogen amendment treatments. Results showed that enriched CO₂ concentration increased canopy WUE, and more N supply led to higher WUE under the increased CO₂. Canopy WUE was significantly lower in well-watered treatments than in drought treatment, but increased with the increased N supply. Elevated CO₂ reduced the apparent recovery fraction of applied N by the plant root system (N_r, defined as the ratio of the increased N uptake to N applied), but increased the NUE and agronomic N efficiency (NAE, defined as the ratio of the increased biomass to N applied). Water limitation and high N application reduced the N_r, NUE and NAE, indicating a poor N efficiency. In addition, there was a close relationship between the root mass ratio and NUE. Canopy WUE was negatively related to the root mass ratio and NUE. Our results indicated that CO₂ enrichment enhanced WUE more at high N application, but increased NUE more when N application was less.     

Effects of elevated CO2 and increased nitrogen deposition on photosynthesis and growth of understory plants in spruce model ecosystems

We studied the effects of atmospheric CO₂ enrichment (280, 420 and 560 l CO₂ l^–1) and increased N deposition (0,30 and 90 kg ha^–1 year^–1) on the spruce-forest understory species Oxalis acetosella, Homogyne alpina and Rubus hirtus. Clones of these species formed the ground cover in nine 0.7 m² model ecosystems with 5-year-old Picea abies trees (leaf area index of approx 2.2). Communities grew on natural forest soil in a simulated montane climate. Independently of N deposition, the rate of light-saturated net photosynthesis of leaves grown and measured at 420 l CO₂ l^–1 was higher in Oxalis and in Homogyne, but was not significantly different in Rubus compared to leaves grown and measured at the pre-industrial CO₂ concentration of 280 l l^–1. Remarkably, further CO₂ enrichment to 560 l l^–1 caused no additional increase of CO₂ uptake. With increasing CO₂ supply concentrations of non-structural carbohydrates in leaves increased and N concentrations decreased in all species, whereas N deposition had no significant effect on these traits. Above-ground biomass and leaf area production were not significantly affected by elevated CO₂ in the more vigorously growing species O. acetosella and R. hirtus, but the slow growing H. alpina produced almost twice as much biomass and 50% more leaf area per plant under 420 l CO₂ l^–1 compared to 280 l l^–1 (again no further stimulation at 560 l l^–1). In contrast, increased N addition stimulated growth in Oxalis and Rubus but had no effect on Homogyne. In Oxalis (only) biomass per plant was positively correlated with microhabitat quantum flux density at low CO₂, but not at high CO₂ indicating carbon saturation. On the other hand, the less shade-tolerant Homogyne profited from CO₂ enrichment at all understory light levels facilitating its spread into more shady micro-habitats under elevated CO₂. These species-specific responses to CO₂ and N deposition will affect community structure. The non-linear responses to elevated CO₂ of several of the traits studied here suggest that the largest responses to rising atmospheric CO₂ are under way now or have already occurred and possible future responses to further increases in CO₂ concentration are likely to be much smaller in these understory species.     

Comparison of the effect of elevated CO2 on an invasive species (Centaurea solstitialis) in monoculture and community settings

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) is likely to change the species composition of plant communities. To investigate whether growth of a highly invasive plant species, Centaurea solstitialis (yellow starthistle), was affected by elevated [CO₂], and whether the success of this species would increase under CO₂ enrichment, I grew the species in serpentine soil microcosms, both as a monoculture and as a component of a grassland community. Centaurea grown in monoculture responded strongly to [CO₂] enrichment of 350 mol mol^–1, increasing aboveground biomass production by 70%, inflorescence production by 74%, and midday photosynthesis by an average of 132%. When grown in competition with common serpentine grassland species, Centaurea responded to CO₂ enrichment with similar but nonsignificant increases (+69% aboveground biomass, +71% inflorescence production), while total aboveground biomass of the polyculture increased by 28%. Centaurea's positive CO₂ response in monoculture and parallel (but non-significant) response in polyculture provoke questions about possible consequences of increasing CO₂ for more typical California grasslands, where the invader already causes major problems.     

Soil-atmosphere exchange of CH4, CO2, NOx,and N2O in the Colorado shortgrass steppe under elevated CO2

In late March 1997, an open-top-chamber (OTC) CO₂ enrichment study was begun in the Colorado shortgrass steppe. The main objectives of the study were to determine the effect of elevated CO₂ (720 mol mol^–1) on plant production, photosynthesis, and water use of this mixed C₃/C₄ plant community, soil nitrogen (N) and carbon (C) cycling and the impact of changes induced by CO₂ on trace gas exchange. From this study, we report here our weekly measurements of CO₂, CH₄, NO_x and N₂O fluxes within control (unchambered), ambient CO₂ and elevated CO₂ OTCs. Soil water and temperature were measured at each flux measurement time from early April 1997, year round, through October 2000. Even though both C₃ and C₄ plant biomass increased under elevated CO₂ and soil moisture content was typically higher than under ambient CO₂ conditions, none of the trace gas fluxes were significantly altered by CO₂ enrichment. Over the 43 month period of observation NO_x and N₂O flux averaged 4.3 and 1.7 in ambient and 4.1 and 1.7 g N m^–2 hr ^–1 in elevated CO₂ OTCs, respectively. NO_x flux was negatively correlated to plant biomass production. Methane oxidation rates averaged –31 and –34 g C m^–2 hr^–1 and ecosystem respiration averaged 43 and 44 mg C m^–2 hr^–1 under ambient and elevated CO₂, respectively, over the same time period.     

Responses of N fluxes and pools to elevated atmospheric CO2 in model forest ecosystems with acidic and calcareous soils

The objectives of this study were to estimate how soil type, elevated N deposition (0.7 vs. 7 g N m^–2y^–1) and tree species influence the potential effects of elevated CO₂ (370 vs. 570 mol CO₂ mol^–1) on N pools and fluxes in forest soils. Model spruce-beech forest ecosystems were established on a nutrient-rich calcareous sand and on a nutrient-poor acidic loam in large open-top chambers. In the fourth year of treatment, we measured N concentrations in the soil solution at different depths, estimated N accumulation by ion exchange resin (IER) bags, and quantified N export in drainage water, denitrification, and net N uptake by trees. Under elevated CO₂, concentrations of N in the soil solution were significantly reduced. In the nutrient-rich calcareous sand, CO₂ enrichment decreased N concentrations in the soil solution at all depths (–45 to –100%). In the nutrient-poor acidic loam, the negative CO₂ effect was restricted to the uppermost 5 cm of the soil. Increasing the N deposition stimulated the negative impact of CO₂ enrichment on soil solution N in the acidic loam at 5 cm depth from –20% at low N inputs to –70% at high N inputs. In the nutrient-rich calcareous sand, N additions did not influence the CO₂ effect on soil solution N. Accumulation of N by IER bags, which were installed under individual trees, was decreased at high CO₂ levels under spruce in both soil types. Under beech, this decrease occurred only in the calcareous sand. N accumulation by IER bags was negatively correlated with current-years foliage biomass, suggesting that the reduction of soil N availability indices was related to a CO₂-induced growth enhancement. However, the net N uptake by trees was not significantly increased by elevated CO₂. Thus, we suppose that the reduced N concentrations in the soil solution at elevated CO₂ concentrations were rather caused by an increased N immobilisation in the soil. Denitrification was not influenced by atmospheric CO₂ concentrations. CO₂ enrichment decreased nitrate leaching in drainage by 65%, which suggests that rising atmospheric CO₂ potentially increases the N retention capacity of forest ecosystems.     

Interrelationships Between Nitrogen Supply and Photosynthetic Parameters in Vicia faba L.

We determined for Vicia faba L the influence of nitrogen uptake and accumulation on the values of photon saturated net photosynthetic rate (P _Nmax), quantum yield efficiency (), intercellular CO₂ concentration (C _i), and carboxylation efficiency (C _e). As leaf nitrogen content (N_L) increased, the converged onto a maximum asymptotic value of 0.0664±0.0049 mol(CO₂) mol(quantum)^–1. Also, as N_L increased the C _i value fell to an asymptotic minimum of 115.80±1.59 mol mol^–1, and C _e converged onto a maximum asymptotic value of 1.645±0.054 mol(CO₂) m^–2 s^–1 Pa^–1 and declined to zero at a N_L-intercept equal to 0.596±0.096 g(N) m^–2. fell to zero for an N_L-intercept of 0.660±0.052 g(N) m^–2. As N_L increased, the value of P _Nmax converged onto a maximum asymptotic value of 33.400±2.563 mol(CO₂) m^–2 s^–1. P _N fell to zero for an N_L-intercept of 0.710±0.035 g(N) m^–2. Under variable daily meteorological conditions the values for N_L, specific leaf area (_L), root mass fraction (R_f), P _Nmax, and remained constant for a given N supply. A monotonic decline in the steady-state value of R_f occurred with increasing N supply. _L increased with increasing N supply or with increasing N_L.     

Interactions of elevated CO2, NH3 and O3 on mycorrhizal infection,gas exchange and N metabolism in saplings of Scots pine

Four-year-old saplings of Scots pine (Pinus sylvestris) (L.) were exposed for 11 weeks in controlled-environment chambers to charcoad-filtered air, or to charcoal-filtered air supplemented with NH₃ (40 g m^–3), O₃ (110 g m^–3 during day/ 40 g m^–3 during night) or NH₃+O₃. All treatments were carried out at ambient (259 L L^–1) and at elevated CO₂ concentration (700 L L^–1). Total tree biomass, mycorrhizal infection, net CO₂ assimilation (P_n), stomatal conductance (g_s), transpiration of the shoots and NH₃ metabolization of the needles were measured. In ambient CO₂ (1) gaseous NH₃ decreased mycorrhizal infection, without significantly affecting tree biomass or N concentration and it enhanced the activity of glutamine synthetase (GS) and glutamate dehydrogenase (GDH) in one-year-old needles; (2) ozone decreased mycorrhizal infection and the acitivity of GS in the needles, while it increased the activity og GDH; (3) exposure to NH₃+O₃ lessened the effects of single exposures to NH₃ and O₃ on reduction of mycorrhizal infection and on increase in GDH activity. Similar lessing effects on mycorrhizal infection as observed in trees exposed to NH₃+O₃ at ambient CO₂, were measured in trees exposed to NH₃+O₃ at elevated CO₂. Exposure to elevated CO₂ without pollutants did not significantly affect any of the parameters studied, except for a decrease in the concentration of soluble proteins in the needles. Elevated CO₂ _NH₃ strongly decreased root branching and mycorrhizal infection and temporarily stimulated P_n and g_s. The exposure to elevated CO₂+NH₃+O₃ also transiently stimulated P_n. The possible mechanisms underlying and integrating these effects are discussed. Elevated CO₂ clearly did not alleviate the negative effects of NH₃ and O₃ mycoorhiral infection. The significant reduction of mycorrhizal infection after exposure to NH₃ or O₃, observed before significant changes in gas exchange or growth occurred, suggest the use of mycorrhizal infection as an early indicator for NH₃ and O₃ induced stress.Abbreviations DW dry weight - FA filtered air - FA_a filtered air at ambient CO₂ - FA_e filtered air at elevated CO₂ - FW fresh weight - GDH glutamate dehydrogenase - GS glutamine synthetase - g_s stomatal conductance - P_n net CO₂ assimilation - RWR root weight ratio - SRL specific root length     

Elevated CO2 alters anatomy,physiology, growth,and reproduction of red mangrove (Rhizophora mangle L.)

Mangroves, woody halophytes restricted to protected tropical coasts, form some of the most productive ecosystems in the world, but their capacity to act as a carbon source or sink under climate change is unknown. Their ability to adjust growth or to function as potential carbon sinks under conditions of rising atmospheric CO₂ during global change may affect global carbon cycling, but as yet has not been investigated experimentally. Halophyte responses to CO₂ doubling may be constrained by the need to use carbon conservatively under water-limited conditions, but data are lacking to issue general predictions. We describe the growth, architecture, biomass allocation, anatomy, and photosynthetic physiology of the predominant neotropical mangrove tree, Rhizophora mangle L., grown solitarily in ambient (350 ll^–1) and double-ambient (700 ll^–1) CO₂ concentrations for over 1 year. Mangrove seedlings exhibited significantly increased biomass, total stem length, branching activity, and total leaf area in elevated CO₂. Enhanced total plant biomass under high CO₂ was associated with higher root:shoot ratios, relative growth rates, and net assimilation rates, but few allometric shifts were attributable to CO₂ treatment independent of plant size. Maximal photosynthetic rates were enhanced among high-CO₂ plants while stomatal conductances were lower, but the magnitude of the treatment difference declined over time, and high-CO₂ seedlings showed a lower P_max at 700 ll^–1 CO₂ than low-CO₂ plants transferred to 700 ll^–1 CO₂: possible evidence of downregulation. The relative thicknesses of leaf cell layers were not affected by treatment. Stomatal density decreased as epidermal cells enlarged in elevated CO₂. Foliar chlorophyll, nitrogen, and sodium concentrations were lower in high CO₂. Mangroves grown in high CO₂ were reproductive after only 1 year of growth (fully 2 years before they typically reproduce in the field), produced aerial roots, and showed extensive lignification of the main stem; hence, elevated CO₂ appeared to accelerate maturation as well as growth. Data from this long-term study suggest that certain mangrove growth characters will change flexibly as atmospheric CO₂ increases, and accord with responses previously shown in Rhizophora apiculata. Such results must be integrated with data from sea-level rise studies to yield predictions of mangrove performance under changing climate.     

Elevated CO2 increases belowground respiration in California grasslands

This study was designed to identify potential effects of elevated CO₂ on belowground respiration (the sum of root and heterotrophic respiration) in field and microcosm ecosystems and on the annual carbon budget. We made three sets of respiration measurements in two CO₂ treatments, i.e., (1) monthly in the sandstone grassland and in microcosms from November 1993 to June 1994; (2) at the annual peak of live biomass (March and April) in the serpentine and sandstone grasslands in 1993 and 1994; and (3) at peak biomass in the microcosms with monocultures of seven species in 1993. To help understand ecosystem carbon cycling, we also made supplementary measurements of belowground respiration monthly in sandstone and serpentine grasslands located within 500 m of the CO₂ experiment site. The seasonal average respiration rate in the sandstone grassland was 2.12 mol m^-2 s^-1 in elevated CO₂, which was 42% higher than the 1.49 mol m^-2 s^-1 measured in ambient CO₂ (P=0.007). Studies of seven individual species in the microcosms indicated that respiration was positively correlated with plant biomass and increased, on average, by 70% with CO₂. Monthly measurements revealed a strong seasonality in belowground respiration, being low (0–0.5 mol CO₂ m^-2 s^-1 in the two grasslands adjacent to the CO₂ site) in the summer dry season and high (2–4 mol CO₂ m^-2 s^-1 in the sandstone grassland and 2–7 mol CO₂ m^-2 s^-1 in the microcosms) during the growing season from the onset of fall rains in November to early spring in April and May. Estimated annual carbon effluxes from the soil were 323 and 440 g C m^-2 year^-1 for the sandstone grasslands in ambient and elevated CO₂. That CO₂-stimulated increase in annual soil carbon efflux is more than twice as big as the increase in aboveground net primary productivity (NPP_a) and approximately 60% of NPP_a in this grassland in the current CO₂ environment. The results of this study suggest that below-ground respiration can dissipate most of the increase in photosynthesis stimulated by elevated CO₂.CIWDPB Publication # 1271     

Modelling the CO2 gas exchange of grassland vegetation from experimental data

Pseudo-cubic spline functions were applied to the two atmospheric CO₂ concentrations of 340 and 600 l·l^–1 CO₂ for describing the average daily CO₂ gas exchange rates of simplifed grassland model ecosystems. Measurements used were from daily means of photon flux density (PhAR), temperature and relative air humidity, phytomass of each day, leaf area indexes, average phenological states of the vegetation (1–15), and water exchange rates of the entire model ecosystems. The functions were validated with, data from the same experimental year. We also succeeded in verifying the functions with the response data from years other than those used for constructing the model.