Review of elevated atmospheric CO2 effects on agro-ecosystems: residue decomposition processes and soil C storage

A series of studies using major crops (cotton [Gossypium hirsutum L.], wheat [Triticum aestivum L.], grain sorghum [Sorghum bicolor (L.) Moench.] and soybean [Glycine max (L.) Merr.]) were reviewed to examine the impact of elevated atmospheric CO₂ on crop residue decomposition within agro-ecosystems. Experiments evaluated utilized plant and soil material collected from CO₂ study sites using Free Air CO₂ Enrichment (FACE) and open top chambers (OTC). A incubation study of FACE residue revealed that CO₂-induced changes in cotton residue composition could alter decomposition processes, with a decrease in N mineralization observed with FACE, which was dependent on plant organ and soil series. Incubation studies utilizing plant material grown in OTC considered CO₂-induced changes in relation to quantity and quality of crop residue for two species, soybean and grain sorghum. As with cotton, N mineralization was reduced with elevated CO₂ in both species, however, difference in both quantity and quality of residue impacted patterns of C mineralization. Over the short-term (14 d), little difference was observed for CO₂ treatments in soybean, but C mineralization was reduced with elevated CO₂ in grain sorghum. For longer incubation periods (60 d), a significant reduction in CO₂-C mineralized per g of residue added was observed with the elevated atmospheric CO₂ treatment in both crop species. Results from incubation studies agreed with those from the OTC field observations for both measurements of short-term CO₂ efflux following spring tillage and the cumulative effect of elevated CO₂ (> 2 years) in this study. Observations from field and laboratory studies indicate that with elevated atmospheric CO₂, the rate of plant residue decomposition may be limited by N and the release of N from decomposing plant material may be slowed. This indicates that understanding N cycling as affected by elevated CO₂ is fundamental to understanding the potential for soil C storage on a global scale. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of elevated atmospheric CO2 in agro-ecosystems on soil carbon storage

Increasing global atmospheric CO₂ concentration has led to concerns regarding its potential effects on the terrestrial environment. Attempts to balance the atmospheric carbon (C) budget have met with a large shortfall in C accounting (≈1.4 × 10¹⁵ g C y^–1) and this has led to the hypothesis that C is being stored in the soil of terrestrial ecosystems. This study examined the effects of CO₂ enrichment on soil C storage in C3 soybean (Glycine max L.) Merr. and C4 grain sorghum (Sorghum bicolor L.) Moench. agro-ecosystems established on a Blanton loamy sand (loamy siliceous, thermic, Grossarenic Paleudults). The study was a split-plot design replicated three times with two crop species (soybean and grain sorghum) as the main plots and two CO₂ concentration (ambient and twice ambient) as subplots using open top field chambers. Carbon isotopic techniques using δ¹³C were used to track the input of new C into the soil system. At the end of two years, shifts in δ¹³C content of soil organic matter carbon were observed to a depth of 30 cm. Calculated new C in soil organic matter with grain sorghum was greater for elevated CO₂ vs. ambient CO₂ (162 and 29 g m^–2, respectively), but with soybean the new C in soil organic matter was less for elevated CO₂ vs. ambient CO₂ (120 and 291 g m^–2, respectively). A significant increase in mineral associated organic C was observed in 1993 which may result in increased soil C storage over the long-term, however, little change in total soil organic C was observed under either plant species. These data indicate that elevated atmospheric CO₂ resulted in changes in soil C dynamics in agro-ecosystems that are crop species dependent.     

Effects of elevated CO2 on the protein concentration of food crops: a meta-analysis

Meta‐analysis techniques were used to examine the effect of elevated atmospheric carbon dioxide [CO₂] on the protein concentrations of major food crops, incorporating 228 experimental observations on barley, rice, wheat, soybean and potato. Each crop had lower protein concentrations when grown at elevated (540–958 μmol mol^?1) compared with ambient (315–400 μmol mol^?1) CO₂. For wheat, barley and rice, the reduction in grain protein concentration was ～10–15% of the value at ambient CO₂. For potato, the reduction in tuber protein concentration was 14%. For soybean, there was a much smaller, although statistically significant reduction of protein concentration of 1.4%. The magnitude of the CO₂ effect on wheat grains was smaller under high soil N conditions than under low soil N. Protein concentrations in potato tubers were reduced more for plants grown at high than at low concentrations of ozone. For soybean, the ozone effect was the reverse, as elevated CO₂ increased the protein concentration of soybean grown at high ozone concentrations. The magnitude of the CO₂ effect also varied depending on experimental methodology. For both wheat and soybean, studies performed in open‐top chambers produced a larger CO₂ effect than those performed using other types of experimental facilities. There was also indication of a possible pot artifact as, for both wheat and soybean, studies performed in open‐top chambers showed a significantly greater CO₂ effect when plants were rooted in pots rather than in the ground. Studies on wheat also showed a greater CO₂ effect when protein concentration was measured in whole grains rather than flour. While the magnitude of the effect of elevated CO₂ varied depending on the experimental procedures, a reduction in protein concentration was consistently found for most crops. These findings suggest that the increasing CO₂ concentrations of the 21st century are likely to decrease the protein concentration of many human plant foods.     

Interactions of tropospheric CO2 and O3 enrichments and moisture variations on microbial biomass and respiration in soil

Soil microbial biomass C (C_mic) is a sensitive indicator of trends in organic matter dynamics in terrestrial ecosystems. This study was conducted to determine the effects of tropospheric CO₂ or O₃ enrichments and moisture variations on total soil organic C (C_org), mineralizable C fraction (C_Min), C_mic, maintenance respiratory (qCO₂) or C_mic death (qD) quotients, and their relationship with basal respiration (BR) rates and field respiration (FR) fluxes in wheat‐soybean agroecosystems. Wheat (Triticum aestivum L.) and soybean (Glycine max. L. Merr) plants were grown to maturity in 3‐m dia open‐top field chambers and exposed to charcoal‐filtered (CF) air at 350 μL CO₂ L^?1; CF air + 150 μL CO₂ L^?1; nonfiltered (NF) air + 35 nL O₃ L^?1; and NF air + 35 nL O₃ L^?1 + 150 μL CO₂ L^?1 at optimum (? 0.05 MPa) and restricted soil moisture (? 1.0 ± 0.05 MPa) regimes. The + 150 μL CO₂ L^?1 additions were 18 h d^?1 and the + 35 nL O₃ L^?1 treatments were 7 h d^?1 from April until late October. While C_org did not vary consistently, C_Min, C_mic and C_mic fractions increased in soils under tropospheric CO₂ enrichment (500 μL CO₂ L^?1) and decreased under high O₃ exposures (55 ± 6 nL O₃ L^?1 for wheat; 60 ± 5 nL O₃ L^?1 for soybean) compared to the CF treatments (25 ± 5 nL O₃ L^?1). The qCO₂ or qD quotients of C_mic were also significantly decreased in soils under high CO₂ but increased under high O₃ exposures compared to the CF control. The BR rates did not vary consistently but they were higher in well‐watered soils. The FR fluxes were lower under high O₃ exposures compared to soils under the CF control. An increase in C_mic or C_mic fractions and decrease in qCO₂ or qD observed under high CO₂ treatment suggest that these soils were acting as C sinks whereas, reductions in C_mic or C_mic fractions and increase in qCO₂ or qD in soils under elevated tropospheric O₃ exposures suggest the soils were serving as a source of CO₂.     

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.     

30 years of free‐air carbon dioxide enrichment (FACE): What have we learned about future crop productivity and its potential for adaptation?

Free‐air CO₂ enrichment (FACE) allows open‐air elevation of [CO₂] without altering the microclimate. Its scale uniquely supports simultaneous study from physiology and yield to soil processes and disease. In 2005 we summarized results of then 28 published observations by meta‐analysis. Subsequent studies have combined FACE with temperature, drought, ozone, and nitrogen treatments. Here, we summarize the results of now almost 250 observations, spanning 14 sites and five continents. Across 186 independent studies of 18 C₃ crops, elevation of [CO₂] by ca. 200 ppm caused a ca. 18% increase in yield under non‐stress conditions. Legumes and root crops showed a greater increase and cereals less. Nitrogen deficiency reduced the average increase to 10%, as did warming by ca. 2°C. Two conclusions of the 2005 analysis were that C₄ crops would not be more productive in elevated [CO₂], except under drought, and that yield responses of C₃ crops were diminished by nitrogen deficiency and wet conditions. Both stand the test of time. Further studies of maize and sorghum showed no yield increase, except in drought, while soybean productivity was negatively affected by early growing season wet conditions. Subsequent study showed reduced levels of nutrients, notably Zn and Fe in most crops, and lower nitrogen and protein in the seeds of non‐leguminous crops. Testing across crop germplasm revealed sufficient variation to maintain nutrient content under rising [CO₂]. A strong correlation of yield response under elevated [CO₂] to genetic yield potential in both rice and soybean was observed. Rice cultivars with the highest yield potential showed a 35% yield increase in elevated [CO₂] compared to an average of 14%. Future FACE experiments have the potential to develop cultivars and management strategies for co‐promoting sustainability and productivity under future elevated [CO₂].     

Soil 13C–15N dynamics in an N2-fixing clover system under long-term exposure to elevated atmospheric CO2

Reduced soil N availability under elevated CO₂ may limit the plant's capacity to increase photosynthesis and thus the potential for increased soil C input. Plant productivity and soil C input should be less constrained by available soil N in an N₂‐fixing system. We studied the effects of Trifolium repens (an N₂‐fixing legume) and Lolium perenne on soil N and C sequestration in response to 9 years of elevated CO₂ under FACE conditions. ¹⁵N‐labeled fertilizer was applied at a rate of 140 and 560 kg N ha^?1 yr^?1 and the CO₂ concentration was increased to 60 Pa pCO₂ using ¹³C‐depleted CO₂. The total soil C content was unaffected by elevated CO₂, species and rate of ¹⁵N fertilization. However, under elevated CO₂, the total amount of newly sequestered soil C was significantly higher under T. repens than under L. perenne. The fraction of fertilizer‐N (f_N) of the total soil N pool was significantly lower under T. repens than under L. perenne. The rate of N fertilization, but not elevated CO₂, had a significant effect on f_N values of the total soil N pool. The fractions of newly sequestered C (f_C) differed strongly among intra‐aggregate soil organic matter fractions, but were unaffected by plant species and the rate of N fertilization. Under elevated CO₂, the ratio of fertilizer‐N per unit of new C decreased under T. repens compared with L. perenne. The L. perenne system sequestered more ¹⁵N fertilizer than T. repens: 179 vs. 101 kg N ha^?1 for the low rate of N fertilization and 393 vs. 319 kg N ha^?1 for the high N‐fertilization rate. As the loss of fertilizer‐¹⁵N contributed to the ¹⁵N‐isotope dilution under T. repens, the input of fixed N into the soil could not be estimated. Although N₂ fixation was an important source of N in the T. repens system, there was no significant increase in total soil C compared with a non‐N₂‐fixing L. perenne system. This suggests that N₂ fixation and the availability of N are not the main factors controlling soil C sequestration in a T. repens system.     

Changes in soil carbon and nitrogen properties and microbial communities in relation to growth of Pinus radiata and Nothofagus fusca trees after 6 years at ambient and elevated atmospheric CO2

Increasing atmospheric CO₂ concentration can influence the growth and chemical composition of many plant species, and thereby affect soil organic matter pools and nutrient fluxes. Here, we examine the effects of ambient (initially 362 μL L^?1) and elevated (654 μL L^?1) CO₂ in open‐top chambers on the growth after 6 years of two temperate evergreen forest species: an exotic, Pinus radiata D. Don, and a native, Nothofagus fusca (Hook. F.) Oerst. (red beech). We also examine associated effects on selected carbon (C) and nitrogen (N) properties in litter and mineral soil, and on microbial properties in rhizosphere and hyphosphere soil. The soil was a weakly developed sand that had a low initial C concentration of about 1.0 g kg^?1 at both 0–100 and 100–300 mm depths; in the N. fusca system, it was initially overlaid with about 50 mm of forest floor litter (predominantly FH material) taken from a Nothofagus forest. A slow‐release fertilizer was added during the early stages of plant growth; subsequent foliage analyses indicated that N was not limiting. After 6 years, stem diameters, foliage N concentrations and C/N ratios of both species were indistinguishable (P>0.10) in the two CO₂ treatments. Although total C contents in mineral soil at 0–100 mm depth had increased significantly (P<0.001) after 6 years growth of P. radiata, averaging 80±0.20 g m^?2 yr^?1, they were not significantly influenced by elevated CO₂. However, CO₂‐C production in litter, and CO₂‐C production, microbial C, and microbial C/N ratios in mineral soil (0–100 mm depth) under P. radiata were significantly higher under elevated than ambient CO₂. CO₂‐C production, microbial C, and numbers of bacteria (but not fungi) were also significantly higher under elevated CO₂ in hyphosphere soil, but not in rhizosphere soil. Under N. fusca, some incorporation of the overlaid litter into the mineral soil had probably occurred; except for CO₂‐C production and microbial C in hyphosphere soil, none of the biochemical properties or microbial counts increased significantly under elevated CO₂. Net mineral‐N production, and generally the potential utilization of different substrates by microbial communities, were not significantly influenced by elevated CO₂ under either tree species. Physiological profiles of the microbial communities did, however, differ significantly between rhizosphere and hyphosphere samples and between samples under P. radiata and N. fusca. Overall, results support the concept that a major effect on soil properties after prolonged exposure of trees to elevated CO₂ is an increase in the amounts, and mineralization rate, of labile organic components.     

Effects of elevated CO2 on the foraging behavior of cotton bollworm, Helicoverpa armigera

Effects of elevated CO2 on the foraging behavior of cotton bollworm Helicoverpa arrnigera Hübner reared on milky grains of spring wheat grown in ambient, 550μL/L and 750μL/L CO2 concentration atmospheres in open-top chambers （OTC） were studied. The results indicated that： （i） elevated CO2 significantly affected both the type and amount of food eaten by H.arrnigera reared on milky grains of ambient CO2-grown wheat were significant higher than those for bollworm larvae reared on wheat grains grown in 550 and 750μL/L CO2 atmospheres; （ii） when bollworm larvae were reared on mixed milky grains from different CO2-grown wheat （food-choice condition）, larval duration increased significantly-pupal weight, adult longevity, and fecundity decreased significantly, comparing with those reared on milky grains of ambient CO2-grown wheat, 550μL/L CO2-grown wheat and 750μL/L CO2-grown wheat respectively; （iii） significant decreases in the contents of fructose and gross protein （GP） and significant increases in the contents of glucose, amylose, total saccharides （TSC）, TSC： GP ratio, free amino acids and soluble protein in the wheat grains with CO2 rising; （iv） and selected-foraging amount/food-choice index of cotton bollworm H.armigera were significantly positive correlated with the contents of fructose and GP of wheat grains, but they had significantly negative relationships with the contents of glucose, amylose, TSC and TSC： GP ratio of wheat grains.     

Decomposition of soil and plant carbon from pasture systems after 9 years of exposure to elevated CO2: impact on C cycling and modeling

Elevated atmospheric CO₂ may alter decomposition rates through changes in plant material quality and through its impact on soil microbial activity. This study examines whether plant material produced under elevated CO₂ decomposes differently from plant material produced under ambient CO₂. Moreover, a long‐term experiment offered a unique opportunity to evaluate assumptions about C cycling under elevated CO₂ made in coupled climate–soil organic matter (SOM) models. Trifolium repens and Lolium perenne plant materials, produced under elevated (60 Pa) and ambient CO₂ at two levels of N fertilizer (140 vs. 560 kg ha^?1 yr^?1), were incubated in soil for 90 days. Soils and plant materials used for the incubation had been exposed to ambient and elevated CO₂ under free air carbon dioxide enrichment conditions and had received the N fertilizer for 9 years. The rate of decomposition of L. perenne and T. repens plant materials was unaffected by elevated atmospheric CO₂ and rate of N fertilization. Increases in L. perenne plant material C : N ratio under elevated CO₂ did not affect decomposition rates of the plant material. If under prolonged elevated CO₂ changes in soil microbial dynamics had occurred, they were not reflected in the rate of decomposition of the plant material. Only soil respiration under L. perenne, with or without incorporation of plant material, from the low‐N fertilization treatment was enhanced after exposure to elevated CO₂. This increase in soil respiration was not reflected in an increase in the microbial biomass of the L. perenne soil. The contribution of old and newly sequestered C to soil respiration, as revealed by the ¹³C‐CO₂ signature, reflected the turnover times of SOM–C pools as described by multipool SOM models. The results do not confirm the assumption of a negative feedback induced in the C cycle following an increase in CO₂, as used in coupled climate–SOM models. Moreover, this study showed no evidence for a positive feedback in the C cycle following additional N fertilization.     

Decomposition of soybean grown under elevated concentrations of CO2 and O3

A critical global climate change issue is how increasing concentrations of atmospheric CO₂ and ground‐level O₃ will affect agricultural productivity. This includes effects on decomposition of residues left in the field and availability of mineral nutrients to subsequent crops. To address questions about decomposition processes, a 2‐year experiment was conducted to determine the chemistry and decomposition rate of aboveground residues of soybean (Glycine max (L.) Merr.) grown under reciprocal combinations of low and high concentrations of CO₂ and O₃ in open‐top field chambers. The CO₂ treatments were ambient (370 μmol mol^?1) and elevated (714 μmol mol^?1) levels (daytime 12 h averages). Ozone treatments were charcoal‐filtered air (21 nmol mol^?1) and nonfiltered air plus 1.5 times ambient O₃ (74 nmol mol^?1) 12 h day^?1. Elevated CO₂ increased aboveground postharvest residue production by 28–56% while elevated O₃ suppressed it by 15–46%. In combination, inhibitory effects of added O₃ on biomass production were largely negated by elevated CO₂. Plant residue chemistry was generally unaffected by elevated CO₂, except for an increase in leaf residue lignin concentration. Leaf residues from the elevated O₃ treatments had lower concentrations of nonstructural carbohydrates, but higher N, fiber, and lignin levels. Chemical composition of petiole, stem, and pod husk residues was only marginally affected by the elevated gas treatments. Treatment effects on plant biomass production, however, influenced the content of chemical constituents on an areal basis. Elevated CO₂ increased the mass per square meter of nonstructural carbohydrates, phenolics, N, cellulose, and lignin by 24–46%. Elevated O₃ decreased the mass per square meter of these constituents by 30–48%, while elevated CO₂ largely ameliorated the added O₃ effect. Carbon mineralization rates of component residues from the elevated gas treatments were not significantly different from the control. However, N immobilization increased in soils containing petiole and stem residues from the elevated CO₂, O₃, and combined gas treatments. Mass loss of decomposing leaf residue from the added O₃ and combined gas treatments was 48% less than the control treatment after 20 weeks, while differences in decomposition of petiole, stem, and husk residues among treatments were minor. Decreased decomposition of leaf residues was correlated with lower starch and higher lignin levels. However, leaf residues only comprised about 20% of the total residue biomass assayed so treatment effects on mass loss of total aboveground residues were relatively small. The primary influence of elevated atmospheric CO₂ and O₃ concentrations on decomposition processes is apt to arise from effects on residue mass input, which is increased by elevated CO₂ and suppressed by O₃.     

Total soil C and N sequestration in a grassland following 10 years of free air CO2 enrichment

Soil C sequestration may mitigate rising levels of atmospheric CO_2. However, it has yet to be determined whether net soil C sequestration occurs in N‐rich grasslands exposed to long‐term elevated CO₂. This study examined whether N‐fertilized grasslands exposed to elevated CO₂ sequestered additional C. For 10 years, Lolium perenne, Trifolium repens, and the mixture of L. perenne/T. repens grasslands were exposed to ambient and elevated CO₂ concentrations (35 and 60 Pa pCO₂). The applied CO₂ was depleted in δ¹³C and the grasslands received low (140 kg ha^?1) and high (560 kg ha^?1) rates of ¹⁵N‐labeled fertilizer. Annually collected soil samples from the top 10 cm of the grassland soils allowed us to follow the sequestration of new C in the surface soil layer. For the first time, we were able to collect dual‐labeled soil samples to a depth of 75 cm after 10 years of elevated CO₂ and determine the total amount of new soil C and N sequestered in the whole soil profile. Elevated CO₂, N‐fertilization rate, and species had no significant effect on total soil C. On average 9.4 Mg new C ha^?1 was sequestered, which corresponds to 26.5% of the total C. The mean residence time of the C present in the 0–10 cm soil depth was calculated at 4.6±1.5 and 3.1±1.1 years for L. perenne and T. repens soil, respectively. After 10 years, total soil N and C in the 0–75 cm soil depth was unaffected by CO₂ concentration, N‐fertilization rate and plant species. The total amount of ¹⁵N‐fertilizer sequestered in the 0–75 cm soil depth was also unaffected by CO₂ concentration, but significantly more ¹⁵N was sequestered in the L. perenne compared with the T. repens swards: 620 vs. 452 kg ha^?1 at the high rate and 234 vs. 133 kg ha^?1 at the low rate of N fertilization. Intermediate values of ¹⁵N recovery were found in the mixture. The fertilizer derived N amounted to 2.8% of total N for the low rate and increased to 8.6% for the high rate of N application. On average, 13.9% of the applied ¹⁵N‐fertilizer was recovered in the 0–75 cm soil depth in soil organic matter in the L. perenne sward, whereas 8.8% was recovered under the T. repens swards, indicating that the N₂‐fixing T. repens system was less effective in sequestering applied N than the non‐N₂‐fixing L. perenne system. Prolonged elevated CO₂ did not lead to an increase in whole soil profile C and N in these fertilized pastures. The potential use of fertilized and regular cut pastures as a net soil C sink under long‐term elevated CO₂ appears to be limited and will likely not significantly contribute to the mitigation of anthropogenic C emissions.     

Smaller than predicted increase in aboveground net primary production and yield of field-grown soybean under fully open-air [CO2] elevation

The Intergovernmental Panel on Climate Change projects that atmospheric [CO₂] will reach 550 ppm by 2050. Numerous assessments of plant response to elevated [CO₂] have been conducted in chambers and enclosures, with only a few studies reporting responses in fully open‐air, field conditions. Reported yields for the world's two major grain crops, wheat and rice, are substantially lower in free‐air CO₂ enrichment (FACE) than predicted from similar elevated [CO₂] experiments within chambers. This discrepancy has major implications for forecasting future global food supply. Globally, the leguminous‐crop soybean (Glycine max (L.) Merr.) is planted on more land than any other dicotyledonous crop. Previous studies have shown that total dry mass production increased on average 37% in response to increasing [CO₂] to approximately 700 ppm, but harvestable yield will increase only 24%. Is this representative of soybean responses under open‐air field conditions? The effects of elevation of [CO₂] to 550 ppm on total production, partitioning and yield of soybean over 3 years are reported. This is the first FACE study of soybean ( http://www.soyface.uiuc.edu ) and the first on crops in the Midwest of North America, one of the major food production regions of the globe. Although increases in both aboveground net primary production (17–18%) and yield (15%) were consistent across three growing seasons and two cultivars, the relative stimulation was less than projected from previous chamber experiments. As in previous studies, partitioning to seed dry mass decreased; however, net production during vegetative growth did not increase and crop maturation was delayed, not accelerated as previously reported. These results suggest that chamber studies may have over‐estimated the stimulatory effect of rising [CO₂], with important implications on global food supply forecasts.     

Root exudation (net efflux of amino acids) may increase rhizodeposition under elevated CO2

Increases in atmospheric CO₂ concentration ([CO₂]) can lead to global climate change and theoretically could enhance carbon (C) deposition in soil, but data on this complex issue are contradictory. One approach for clarifying the diverse forces influencing plant‐derived C in the rhizosphere involves defining how elevated [CO₂] alters the fundamental process of C transfer from plant roots to the soil. We examine here how a step increase in [CO₂] affects the innate influx and efflux components of root exudation in axenic plants, as one foundation for understanding how climate change may affect rhizodeposition. Increasing [CO₂] from 425 to 850 μmol mol^?1 during short‐term trials enhanced shoot and root dry weight (P<0.01) of annual rye grass (Lolium multiflorum Lam.) and medic (Medicago truncatula L.) but had no effect on growth of maize (Zea mays L.). Root amino‐acid flux in the same plants changed only in maize, which increased the efflux rate (nmol g root fresh weight^?1 h^?1) of six amino acids (arginine, alanine, proline, tyrosine, lysine and leucine) significantly (P<0.05) under elevated [CO₂]. None of the three plant species altered the steady‐state concentration of 16 amino acids released into a hydroponic solution with changing [CO₂], apparently because amino‐acid influx rates, measured at 2.5 μm , consistently exceeded efflux rates. Indeed, plants recovered amino acids at rates 94–374% higher than they were lost from roots regardless of [CO₂]. These results indicate that, in theory, any effect of [CO₂] doubling on amino‐acid efflux can be offset by innately higher rates of influx. In practice, however, higher rates of amino‐acid cycling (i.e., efflux+influx) for each root segment (in C₄ maize) or from more root tissue (in the two C₃ species) should increase root exudation by plants exposed to elevated [CO₂] as additional amino acids would be adsorbed to soil particles or be taken up by soil microorganisms.     

Elevated carbon dioxide and irrigation effects on water stable aggregates in a Sorghum field: a possible role for arbuscular mycorrhizal fungi

While soil biota and processes are becoming increasingly appreciated as important parameters for consideration in global change studies, the fundamental characteristic of soil structure is a neglected area of research. In a sorghum [Sorghum bicolor (L.) Moench] field experiment in which CO₂[supplied using free‐air CO₂ enrichment (FACE) technology] was crossed factorially with an irrigation treatment, soil aggregate (1–2 mm) water stability increased in response to elevated CO₂. Aggregate water stability was increased by 40% and 20% in response to CO₂, at ample and limited water supply treatments, respectively. Soil hyphal lengths of arbuscular mycorrhizal fungi (AMF) increased strongly (with a threefold increase in the dry treatment) in response to CO₂, and the concentrations of one fraction (easily extractable glomalin, EEG) of the AMF‐produced protein glomalin were also increased. Two fractions of glomalin, and AMF hyphal lengths were all positively correlated with soil aggregate water stability. The present results further support the hypothesis that AMF can become important in global change scenarios. Although in this field study a causal relationship between hyphal length, glomalin and aggregate stability cannot be demonstrated, the present data do suggest that AMF could mediate changes in soil structure under elevated CO₂. This could be of great importance in agricultural systems threatened by erosional soil loss.     

Net soil carbon input under ambient and elevated CO2 concentrations: isotopic evidence after 4 years

Elevation of atmospheric CO₂ concentration is predicted to increase net primary production, which could lead to additional C sequestration in terrestrial ecosystems. Soil C input was determined under ambient and Free Atmospheric Carbon dioxide Enrichment (FACE) conditions for Lolium perenne L. and Trifolium repens L. grown for four years in a sandy‐loam soil. The ¹³C content of the soil organic matter C had been increased by 5‰ compared to the native soil by prior cropping to corn (Zea mays) for > 20 years. Both species received low or high amounts of N fertilizer in separate plots. The total accumulated above‐ground biomass produced by L. perenne during the 4‐year period was strongly dependent on the amount of N fertilizer applied but did not respond to increased CO₂. In contrast, the total accumulated above‐ground biomass of T. repens doubled under elevated CO₂ but remained independent of N fertilizer rate. The C:N ratio of above‐ground biomass for both species increased under elevated CO₂ whereas only the C:N ratio of L. perenne roots increased under elevated CO₂. Root biomass of L. perenne doubled under elevated CO₂ and again under high N fertilization. Total soil C was unaffected by CO₂ treatment but dependent on species. After 4 years and for both crops, the fraction of new C (F‐value) under ambient conditions was higher (P= 0.076) than under FACE conditions: 0.43 vs. 0.38. Soil under L. perenne showed an increase in total soil organic matter whereas N fertilization or elevated CO₂ had no effect on total soil organic matter content for both systems. The net amount of C sequestered in 4 years was unaffected by the CO₂ concentration (overall average of 8.5 g C kg^?1 soil). There was a significant species effect and more new C was sequestered under highly fertilized L. perenne. The amount of new C sequestered in the soil was primarily dependent on plant species and the response of root biomass to CO₂ and N fertilization. Therefore, in this FACE study net soil C sequestration was largely depended on how the species responded to N rather than to elevated CO₂.     

Sequestration and turnover of bacterial- and fungal-derived carbon in a temperate grassland soil under long-term elevated atmospheric pCO2

Temperate grasslands contribute about 20% to the global C budget. Elevation of atmospheric CO₂ concentration (pCO₂) could lead to additional C sequestration into these ecosystems. Microbial‐derived C in the soil comprising about 1–5% of total soil organic carbon may be an important ‘pool’ for long‐term storage of C under future increased atmospheric CO₂ concentrations. In our study, the impact of elevated pCO₂ on bacterial‐ and fungal‐derived C in the soil of Lolium perenne pastures was investigated under free air carbon dioxide enrichment (FACE) conditions. For 7 years, L. perenne swards were exposed to ambient and elevated pCO₂ (36 and 60 Pa pCO₂, respectively). The additional CO₂ in the FACE plots was depleted in ¹³C compared with ambient plots, so that ‘new’ (<7 years) C inputs in the form of microbial‐derived residues could be determined by means of stable C isotope analysis. Amino sugars in soil are reliable organic biomarkers for indicating the presence of microbial‐derived residues, with particular amino sugars indicative of either bacterial or fungal origin. It is assumed that amino sugars are stabilized to a significant extent in soil, and so may play an important role in long‐term C storage. In our study, we were also able to discriminate between ‘old’ (> 7 years) and ‘new’ microbial‐derived C using compound‐specific δ¹³C analysis of individual amino sugars. This new tool was very useful in investigating the potential for C storage in microbial‐derived residues and the turnover of this C in soil under increased atmospheric pCO₂. The ¹³C signature of individual amino sugars varied between ?17.4‰ and ?39.6‰, and was up to 11.5% depleted in ¹³C in the FACE plots when compared with the bulk δ¹³C value of the native C3 L. perenne soil. New amino sugars in the bulk soil contributed up to 16% to the overall amino sugar pool after the first year and between 62% and 125% after 7 years of exposure to elevated pCO₂. Amounts of new glucosamine increased by the greatest amount (16–125%) during the experiment, followed by mannosamine (?9% to 107%), muramic acid (?11% to 97%), and galactosamine (15–62%). Proportions of new amino sugars in particle size fractions varied between 38% for muramic acid in the clay fraction and 100% for glucosamine and galactosamine in the coarse sand fraction. Summarizing, during the 7‐year period, amino sugars constituted only between 0.9% and 1.6% of the total SOC content. Therefore, their absolute significance for long‐term C sequestration is limited. Additionally new amino sugars were only sequestered in the silt fraction upon elevated pCO₂ exposure while amino sugar concentrations in the clay fraction decreased. Overall, amino sugar concentrations in bulk soil did not change significantly upon exposure to elevated pCO₂. The calculated mean residence time of amino sugars was surprisingly low varying between 6 and 90 years in the bulk soil, and between 3 and 30 years in the particle size fractions, representing soil organic matter pools with different but relatively low turnover times. Therefore, compound‐specific δ¹³C analysis of individual amino sugars clearly revealed a high amino sugar turnover despite more or less constant amino sugar concentrations over a 7 years period of exposure to elevated pCO₂.     

Above- and below-ground responses of C3–C4 species mixtures to elevated CO2 and soil water availability

We evaluated the influences of CO₂[Control, ～ 370 µ mol mol ^{? 1}; 200 µ mol mol ^{? 1} above ambient applied by free‐air CO₂ enrichment (FACE)] and soil water (Wet, Dry) on above‐ and below‐ground responses of C₃ (cotton, Gossypium hirsutum) and C₄ (sorghum, Sorghum bicolor) plants in monocultures and two density mixtures. In monocultures, CO₂ enrichment increased height, leaf area, above‐ground biomass and reproductive output of cotton, but not sorghum, and was independent of soil water treatment. In mixtures, cotton, but not sorghum, above‐ground biomass and height were generally reduced compared to monocultures, across both CO₂ and soil water treatments. Density did not affect individual plant responses of either cotton or sorghum across the other treatments. Total (cotton + sorghum) leaf area and above‐ground biomass in low‐density mixtures were similar between CO₂ treatments, but increased by 17–21% with FACE in high‐density mixtures, due to a 121% enhancement of cotton leaf area and a 276% increase in biomass under the FACE treatment. Total root biomass in the upper 1.2 m of the soil was not influenced by CO₂ or by soil water in monoculture or mixtures; however, under dry conditions we observed significantly more roots at lower soil depths ( > 45 cm). Sorghum roots comprised 81–85% of the total roots in the low‐density mixture and 58–73% in the high‐density mixture. CO₂‐enrichment partly offset negative effects of interspecific competition on cotton in both low‐ and high‐density mixtures by increasing above‐ground biomass, with a greater relative increase in the high‐density mixture. As a consequence, CO₂‐enrichment increased total above‐ground yield of the mixture at high density. Individual plant responses to CO₂ enrichment in global change models that evaluate mixed plant communities should be adjusted to incorporate feedbacks for interspecific competition. Future field studies in natural ecosystems should address the role that a CO₂‐mediated increase in C₃ growth may have on subsequent vegetation change.     

Nitrogen mineralization and potential nitrification at different depths in acid forest soils

Yield decline of cereals grown in monoculture may be alleviated with alternative crop management strategies. Crop rotation and optimized tillage and fertilizer management can contribute to more sustainable food and fiber production in the long-term by increasing diversity, maintaining soil organic matter (SOM), and reducing adverse effects of excessive N application on water quality. We investigated the effects of crop sequence, tillage, and N fertilization on long-term grain production on an alluvial, silty clay loam soil in southcentral Texas. Crop sequences consisted of monoculture sorghum (Sorghum bicolor (L.) Moench,) wheat (Triticum aestivum L.), and soybean (Glycine max (L.) Merr), wheat/soybean double-crop, and rotation of sorghum with wheat/soybean. Grain yields tended to be lower with no tillage (NT) than with conventional tillage (CT) early in the study and became more similar after 11 years. Nitrogen fertilizer required to produce 95% to maximum sorghum yield was similar for monoculture and rotation upon initiation of the experiment and averaged 16 and 11 mg N g^-1 grain with NT and CT, respectively. After 11 years, however, the N fertilizer requirement became similar for both tillage regimes, but was greater in monoculture (17 mg N g^-1 grain) than in rotation (12 mg N g^-1 grain). Crop sequences with double-cropping resulted in greater land use efficiency because similar or lower amounts of N fertilizer were required to produce equivalent grain than with less intensive monoculture systems. These more intensive crop sequences produced more stover with higher N quality primarily due to the inclusion of soybean in the rotation. Large quantities of stover that remained on the soil surface with NT led to greater SOM content, which increased the internal cycling of nutrients in this soil. In southcentral Texas, where rainfall averages nearly 1000 mm yr^-1, more intensive cropping of sorghum, wheat, and soybean with moderate N fertilization using reduced tillage can increase grain production and potentially decrease N losses to the environment by cycling more N into the crop-SOM system.     

Effect of elevated [CO2] on stem wood properties of mature Norway spruce grown at different soil nutrient availability

The objective of the present study was to investigate the interactive effects of elevated [CO₂] and soil nutrient availability on secondary xylem structure and chemical composition of 41‐year‐old Norway spruce (Picea abies (L.) Karst.) trees. The nonfertilized and irrigated‐fertilized trees were, for 3 years, continuously exposed to elevated [CO₂] in whole‐tree chambers. Elevated [CO₂] decreased concentrations of soluble sugars, acid‐soluble lignin and nitrogen in stem wood, but the effects were not consistent between sampling height and/or fertilization. The effect of 2*ambient [CO₂] on wood structure depended on the exposure year and/or fertilization. Radial lumen diameter decreased and annual ring width increased in the second year of exposure (1999) in elevated [CO₂]. In the latter, the CO₂ effect was significant only in the nonfertilized trees. Stem wood chemistry and structure were significantly affected by fertilization. Fertilization increased the concentrations of nitrogen and gravimetric lignin, annual ring width, and radial lumen diameter. Fertilization decreased C/N ratio, mean ring density, earlywood density, latewood density, cell wall thickness, cell wall index, and latewood percentage. We conclude that elevated [CO₂] had only minor effects on wood properties while fertilization had more marked effects and thus may affect ecosystem processes and suitability of wood for different end‐use purposes.