Carbon and water fluxes in a calcareous grassland under elevated CO2

1. As part of a long-term study of the effects of elevated CO₂ on biodiversity and ecosystem function in a calcareous grassland, we measured ecosystem carbon dioxide and water-vapour fluxes over 24-h periods during the 1994 and 1995 growing seasons. Data were used to derive CO₂ and H₂O gas-exchange response functions to quantum flux density (QFD).
2. The relative increase in net ecosystem CO₂ flux (NEC) owing to CO₂ enrichment increased as QFD rose. Daytime NEC at high QFD under elevated CO₂ increased by 25% to 60%, with the greatest increases in the spring and after mowing in June when above-ground biomass was lowest. There was much less stimulation of NEC in early June and again in October when the canopy was fully developed. Night-time NEC was not significantly altered under elevated CO₂.
3. Short-term reversal of CO₂ concentrations between treatments after two seasons of CO₂ exposure provided evidence for a 50% downward adjustment of NEC expressed per unit above-ground plant dry weight. However, when expressed on a land area basis, this difference disappeared because of a c. 20% increase in above-ground biomass under elevated CO₂.
4. Ecosystem evapotranspiration (ET) was not significantly altered by elevated CO₂ when averaged over all measurement dates and positions. However, ET was reduced 3–18% at high QFD in plots at the top of the slope at our study site. In summary, CO₂ enrichment resulted in a large stimulation of ecosystem CO₂ capture, especially during periods of a large demand of carbon in relationship to its supply, and resulted in a relatively small and variable effect on ecosystem water consumption.     

Water savings in mature deciduous forest trees under elevated CO2

Stomatal conductance of plants exposed to elevated CO₂ is often reduced. Whether this leads to water savings in tall forest‐trees under future CO₂ concentrations is largely unknown but could have significant implications for climate and hydrology. We used three different sets of measurements (sap flow, soil moisture and canopy temperature) to quantify potential water savings under elevated CO₂ in a ca. 35 m tall, ca. 100 years old mixed deciduous forest. Part of the forest canopy was exposed to 540 ppm CO₂ during daylight hours using free air CO₂ enrichment (FACE) and the Swiss Canopy Crane (SCC). Across species and a wide range of weather conditions, sap flow was reduced by 14% in trees subjected to elevated CO₂, yielding ca. 10% reduction in evapotranspiration. This signal is likely to diminish as atmospheric feedback through reduced moistening of the air comes into play at landscape scale. Vapour pressure deficit (VPD)‐sap flow response curves show that the CO₂ effect is greatest at low VPD, and that sap flow saturation tends to occur at lower VPD in CO₂‐treated trees. Matching stomatal response data, the CO₂ effect was largely produced by Carpinus and Fagus, with Quercus contributing little. In line with these findings, soil moisture at 10 cm depth decreased at a slower rate under high‐CO₂ trees than under control trees during rainless periods, with a reversal of this trend during prolonged drought when CO₂‐treated trees take advantage from initial water savings. High‐resolution thermal images taken at different heights above the forest canopy did detect reduced water loss through altered energy balance only at <5 m distance (0.44 K leaf warming of CO₂‐treated Fagus trees). Short discontinuations of CO₂ supply during morning hours had no measurable canopy temperature effects, most likely because the stomatal effects were small compared with the aerodynamic constraints in these dense, broad‐leaved canopies. Hence, on a seasonal basis, these data suggest a <10% reduction in water consumption in this type of forest when the atmosphere reaches 540% ppm CO₂.     

Stomatal sensitivity to vapour pressure difference over a subambient to elevated CO2 gradient in a C3/C4 grassland

In the present study the response of stomatal conductance (g_s) to increasing leaf‐to‐air vapour pressure difference (D) in early season C₃ (Bromus japonicus) and late season C₄ (Bothriochloa ischaemum) grasses grown in the field across a range of CO₂ (200–550 µmol mol^?1) was examined. Stomatal sensitivity to D was calculated as the slope of the response of g_s to the natural log of externally manipulated D (dg_s/dlnD). Increasing D and CO₂ significantly reduced g_s in both species. Increasing CO₂ caused a significant decrease in stomatal sensitivity to D in Br. japonicus, but not in Bo. ischaemum. The decrease in stomatal sensitivity to D at high CO₂ for Br. japonicus fit theoretical expectations of a hydraulic model of stomatal regulation, in which g_s varies to maintain constant transpiration and leaf water potential. The weaker stomatal sensitivity to D in Bo. ischaemum suggested that stomatal regulation of leaf water potential was poor in this species, or that non‐hydraulic signals influenced guard cell behaviour. Photosynthesis (A) declined with increasing D in both species, but analyses of the ratio of intercellular to atmospheric CO₂ (C_i/C_a) suggested that stomatal limitation of A occurred only in Br. japonicus. Rising CO₂ had the greatest effect on g_s and A in Br. japonicus at low D. In contrast, the strength of stomatal and photosynthetic responses to CO₂ were not affected by D in Bo. ischaemum. Carbon and water dynamics in this grassland are dominated by a seasonal transition from C₃ to C₄ photosynthesis. Interspecific variation in the response of g_s to D therefore has implications for predicting seasonal ecosystem responses to CO₂.     

N2 fixation by Acacia species increases under elevated atmospheric CO2

In the present study the effect of elevated CO₂ on growth and nitrogen fixation of seven Australian Acacia species was investigated. Two species from semi‐arid environments in central Australia (Acacia aneura and A. tetragonophylla) and five species from temperate south‐eastern Australia (Acacia irrorata, A. mearnsii, A. dealbata, A. implexa and A. melanoxylon) were grown for up to 148 d in controlled greenhouse conditions at either ambient (350 µmol mol^?1) or elevated (700 µmol mol^?1) CO₂ concentrations. After establishment of nodules, the plants were completely dependent on symbiotic nitrogen fixation. Six out of seven species had greater relative growth rates and lower whole plant nitrogen concentrations under elevated versus normal CO₂. Enhanced growth resulted in an increase in the amount of nitrogen fixed symbiotically for five of the species. In general, this was the consequence of lower whole‐plant nitrogen concentrations, which equate to a larger plant and greater nodule mass for a given amount of nitrogen. Since the average amount of nitrogen fixed per unit nodule mass was unaltered by atmospheric CO₂, more nitrogen could be fixed for a given amount of plant nitrogen. For three of the species, elevated CO₂ increased the rate of nitrogen fixation per unit nodule mass and time, but this was completely offset by a reduction in nodule mass per unit plant mass.     

Eddy-correlation system for long-term monitoring of fluxes of heat, water vapour and CO2

An eddy-correlation system is presented that was designed with special focus on long-term measurements of turbulent fluxes in the atmospheric boundary layer. It consists of a SOLENT sonic anemometer, a fast temperature sensor, and a LI-COR LI 6262 closed-path infrared gas analyser. The use of a fast temperature sensor turned out to be necessary because of errors in the sound virtual temperature measured by the sonic anemometer at high wind speeds. The components are combined with special attention paid to protection against lightning and other environmental stresses. The data acquisition program SOLCOM runs on standalone systems or in a network environment and performs ‘quasi on-line’ data processing, on-line graphical display of single data and fluxes, and on-line correction of the raw data. Raw data can be stored continuously on DAT tapes. All data handling can be done by remote access, thus only a minimum amount of m situ maintenance is required. Power spectra of vertical and longitudinal wind speed, air temperature, air humidity and carbon dioxide concentration showed to follow the -2/3 law quite well. There was some noise in the high frequency range of the carbon dioxide spectrum. However, the corresponding cross spectra with the vertical wind component showed less deviation from a straight line in the high frequency range. The sum of convective heat fluxes and soil heat flux showed good agreement with the measured net radiation for several months and it was concluded that the system described here constitute a good platform for long-term flux measurements over forest.     

Consequences of wildfire on ecosystem CO2 and water vapour fluxes in the Great Basin

Increased fire frequency in the Great Basin of North America's intermountain West has led to large‐scale conversion of native sagebrush (Artemisia tridentata Nutt.) communities to postfire successional communities dominated by native and non‐native annual species during the last century. The consequences of this conversion for basic ecosystem functions, however, are poorly understood. We measured net ecosystem CO₂ exchange (NEE) and evapotranspiration (ET) during the first two dry years after wildfire using a 4‐m diameter (16.4 m³) translucent static chamber (dome), and found that both NEE and ET were higher in a postfire successional ecosystem (?0.9–2.6 µ mol CO₂ m^?2 s^?1 and 0.0–1.0 mmol H₂O m^?2 s^?2, respectively) than in an adjacent intact sagebrush ecosystem (?1.2–2.3 µ mol CO₂ m^?2 s^?1 and ?0.1–0.8 mmol H₂O m^?2 s^?2, respectively) during relatively moist periods. Higher NEE in the postfire ecosystem appears to be due to lower rates of above‐ground plant respiration while higher ET appears to be caused by higher surface soil temperatures and increased soil water recharge after rains. These patterns disappeared or were reversed, however, when the conditions were drier. Daily net ecosystem productivity (NEP; g C m^?2 d^?1), derived from multiple linear regressions of measured fluxes with continuously measured climate variables, was very small (close to zero) throughout most of the year. The wintertime was an exception in the intact sagebrush ecosystem with C losses exceeding C gains leading to negative NEP while C balance of the postfire ecosystem remained near zero. Taken together, our results indicate that wildfire‐induced conversion of native sagebrush steppe to ecosystems dominated by herbaceous annual species may have little effect on C balance during relatively dry years (except in winter months) but may stimulate water loss immediately following fires.     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

Response of wheat canopy CO2 and water gas-exchange to soil water content under ambient and elevated CO2

The nature of the interaction between drought and elevated CO₂ partial pressure (pC_a) is critically important for the effects of global change on crops. Some crop models assume that the relative responses of transpiration and photosynthesis to soil water deficit are unaltered by elevated pC_a, while others predict decreased sensitivity to drought at elevated pC_a. These assumptions were tested by measuring canopy photosynthesis and transpiration in spring wheat (cv. Minaret) stands grown in boxes with 100 L rooting volume. Plants were grown under controlled environments with constant light (300 µmol m^?2 s^?1) at ambient (36 Pa) or elevated (68 Pa) pC_a and were well watered throughout growth or had a controlled decline in soil water starting at ear emergence. Drought decreased final aboveground biomass (?15%) and grain yield (?19%) while elevated pC_a increased biomass (+24%) and grain yield (+29%) and there was no significant interaction. Elevated pC_a increased canopy photosynthesis by 15% on average for both water regimes and increased dark respiration per unit ground area in well‐watered plants, but not drought‐grown ones. Canopy transpiration and photosynthesis were decreased in drought‐grown plants relative to well‐watered plants after about 20–25 days from the start of the drought. Elevated pC_a decreased transpiration only slightly during drought, but canopy photosynthesis continued to be stimulated so that net growth per unit water transpired increased by 21%. The effect of drought on canopy photosynthesis was not the consequence of a loss of photosynthetic capacity initially, as photosynthesis continued to be stimulated proportionately by a fixed increase in irradiance. Drought began to decrease canopy transpiration below a relative plant‐available soil water content of 0.6 and canopy photosynthesis and growth below 0.4. The shape of these responses were unaffected by pC_a, supporting the simple assumption used in some models that they are independent of pC_a.     

18O composition of CO2 and H2O ecosystem pools and fluxes in a tallgrass prairie: Simulations and comparisons to measurements

In this paper we describe measurements and modeling of ¹⁸O in CO₂ and H₂O pools and fluxes at a tallgrass prairie site in Oklahoma. We present measurements of the δ¹⁸O value of leaf water, depth‐resolved soil water, atmospheric water vapor, and Keeling plot δ¹⁸O intercepts for net soil‐surface CO₂ and ecosystem CO₂ and H₂O fluxes during three periods of the 2000 growing season. Daytime discrimination against C¹⁸OO, as calculated from measured above‐canopy CO₂ and δ¹⁸O gradients, is also presented. To interpret the isotope measurements, we applied an integrated land‐surface and isotope model (ISOLSM) that simulates ecosystem H₂¹⁸O and C¹⁸OO stocks and fluxes. ISOLSM accurately predicted the measured isotopic composition of ecosystem water pools and the δ¹⁸O value of net ecosystem CO₂ and H₂O fluxes. Simulations indicate that incomplete equilibration between CO₂ and H₂O within C₄ plant leaves can have a substantial impact on ecosystem discrimination. Diurnal variations in the δ¹⁸O value of above‐canopy vapor had a small impact on the predicted δ¹⁸O value of ecosystem water pools, although sustained differences had a large impact. Diurnal variations in the δ¹⁸O value of above‐canopy CO₂ substantially affected the predicted ecosystem discrimination. Leaves dominate the ecosystem ¹⁸O‐isoflux in CO₂ during the growing season, while the soil contribution is relatively small and less variable. However, interpreting daytime measurements of ecosystem C¹⁸OO fluxes requires accurate predictions of both soil and leaf ¹⁸O‐isofluxes.     

Reduced water repellency of a grassland soil under elevated atmospheric CO2

Water repellency is a widespread characteristic of soils that can modify soil moisture content and distribution and is implicated in important processes such as aggregation and carbon sequestration. Repellency arises as a consequence of organic matter inputs; as elevated atmospheric CO₂ is known to modify such inputs, we tested the repellency of a grassland soil after 5 years of exposure to elevated CO₂ in a free air carbon dioxide enrichment experiment. Using a water droplet penetration time test, we found a significant reduction in repellency at elevated CO₂ in samples at field moisture content. As many of the processes potentially influenced by repellency have been shown to be modified at elevated CO₂ (e.g. soil aggregation, C sequestration, recruitment from seed), we suggest that further exploration of this phenomenon could enhance our understanding of CO₂ effects on ecosystem function. The mechanism responsible for the change in repellency has not been identified.     

The input and fate of new C in two forest soils under elevated CO2

The aim of this study was to estimate (i) the influence of different soil types on the net input of new C into soils under CO₂ enrichment and (ii) the stability and fate of these new C inputs in soils. We exposed young beech–spruce model ecosystems on an acidic loam and calcareous sand for 4 years to elevated CO₂. The added CO₂ was depleted in ¹³C, allowing to trace new C inputs in the plant–soil system. We measured CO₂‐derived new C in soil C pools fractionated into particle sizes and monitored respiration as well as leaching of this new C during incubation for 1 year. Soil type played a crucial role in the partitioning of C. The net input of new C into soils under elevated CO₂ was about 75% greater in the acidic loam than in the calcareous sand, despite a 100% and a 45% greater above‐ and below‐ground biomass on the calcareous sand. This was most likely caused by a higher turnover of C in the calcareous sand as indicated by 30% higher losses of new C from the calcareous sand than from the acidic loam during incubation. Therefore, soil properties determining stabilization of soil C were apparently more important for the accumulation of C in soils than tree productivity. Soil fractionation revealed that about 60% of the CO₂‐derived new soil C was incorporated into sand fractions. Low natural ¹³C abundance and wide C/N ratios show that sand fractions comprise little decomposed organic matter. Consistently, incubation indicated that new soil C was preferentially respired as CO₂. During the first month, evolved CO₂ consisted to 40–55% of new C, whereas the fraction of new C in bulk soil C was 15–23% only. Leaching of DOC accounted for 8–23% of the total losses of new soil C. The overall effects of CO₂ enrichment on soil C were small in both soils, although tree growth increased significantly on the calcareous sand. Our results suggest that the potential of soils for C sequestration is limited, because only a small fraction of new C inputs into soils will become long‐term soil C.     

The impact of elevated CO2 on yield loss from a C3 and C4 weed in field-grown soybean

Soybean (Glycine max) was grown at ambient and enhanced carbon dioxide (CO₂, + 250 μL L^?1 above ambient) with and without the presence of a C3 weed (lambsquarters, Chenopodium album L.) and a C4 weed (redroot pigweed, Amaranthus retroflexus L.), in order to evaluate the impact of rising atmospheric carbon dioxide concentration [CO₂] on crop production losses due to weeds. Weeds of a given species were sown at a density of two per metre of row. A significant reduction in soybean seed yield was observed with either weed species relative to the weed‐free control at either [CO₂]. However, for lambsquarters the reduction in soybean seed yield relative to the weed‐free condition increased from 28 to 39% as CO₂ increased, with a 65% increase in the average dry weight of lambsquarters at enhanced [CO₂]. Conversely, for pigweed, soybean seed yield losses diminished with increasing [CO₂] from 45 to 30%, with no change in the average dry weight of pigweed. In a weed‐free environment, elevated [CO₂] resulted in a significant increase in vegetative dry weight and seed yield at maturity for soybean (33 and 24%, respectively) compared to the ambient CO₂ condition. Interestingly, the presence of either weed negated the ability of soybean to respond either vegetatively or reproductively to enhanced [CO₂]. Results from this experiment suggest: (i) that rising [CO₂] could alter current yield losses associated with competition from weeds; and (ii) that weed control will be crucial in realizing any potential increase in economic yield of agronomic crops such as soybean as atmospheric [CO₂] increases.     

Responses of a C3 and a C4 perennial grass to elevated CO2 and temperature under different water regimes

An experiment was carried out to determine the effects of elevated CO₂, elevated temperatures, and altered water regimes in native shortgrass steppe. Intact soil cores dominated by Bouteloua gracilis, a C₄ perennial grass, or Pascopyrum smithii, a C₃ perennial grass, were placed in growth chambers with 350 or 700 μL L^?1 atmospheric CO₂, and under either normal or elevated temperatures. The normal regime mimicked field patterns of diurnal and seasonal temperatures, and the high-temperature regime was 4 °C warmer. Water was supplied at three different levels in a seasonal pattern similar to that observed in the field. Total biomass after two growing seasons was 19% greater under elevated CO₂, with no significant difference between the C₃ and C₄ grass. The effect of elevated CO₂ on biomass was greatest at the intermediate water level. The positive effect of elevated CO₂ on shoot biomass was greater at normal temperatures in B. gracilis, and greater at elevated temperatures in P. smithii. Neither root-to-shoot ratio nor production of seed heads was affected by elevated CO₂. Plant tissue N and soil inorganic N concentrations were lower under elevated Co₂, but no more so in the C₃ than the C₄ plant. Elevated CO₂ appeared to increase plant N limitation, but there was no strong evidence for an increase in N limitation or a decrease in the size of the CO₂ effect from the first to the second growing season. Autumn samples of large roots plus crowns, the perennial organs, had 11% greater total N under elevated CO₂, in spite of greater N limitation.