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.     

森林土壤融化期异养呼吸和微生物碳变化特征

采用室内土柱培养的方法,研究在不同湿度(55%和80%WFPS,土壤充水孔隙度)和不同氮素供给(NH_4Cl和KNO_3,4.5 g N/m~2)条件下,外源碳添加(葡萄糖,6.4 g C/m~2)对温带成熟阔叶红松混交林和次生白桦林土壤融化过程微生物呼吸和微生物碳的激发效应。结果表明:在整个融化培养期间,次生白桦林土壤对照CO_2累积排放量显著高于阔叶红松混交林土壤。随着土壤湿度的增加,次生白桦林土壤对照CO_2累积排放量和微生物代谢熵(q_(CO_2))显著降低,而阔叶红松混交林土壤两者显著地增加(P0.05)。两种林分土壤由葡萄糖(Glu)引起的CO_2累积排放量(9.61—13.49 g C/m~2)显著大于实验施加的葡萄糖含碳量(6.4g C/m~2),同时由Glu引起的土壤微生物碳增量为3.65—27.18 g C/m~2,而施加Glu对土壤DOC含量影响较小。因此,这种由施加Glu引起的额外碳释放可能来源于土壤固有有机碳分解。融化培养结束时,阔叶红松混交林土壤未施氮处理由Glu引起的CO_2累积排放量在两种湿度条件下均显著大于次生白桦林土壤(P0.001);随着湿度的增加,两种林分土壤Glu引起的CO_2累积排放量显著增大(P0.001)。单施KNO_3显著地增加两种湿度的次生白桦林土壤Glu引起的CO_2累积排放量(P0.01)。单施KNO_3显著地增加了两种湿度次生白桦林土壤Glu引起的微生物碳(P0.001),单施NH_4Cl显著地增加低湿度阔叶红松混交林土壤Glu引起的微生物碳(P0.001)。结合前期报道的未冻结实验结果,发现冻结过程显著地影响外源Glu对温带森林土壤微生物呼吸和微生物碳的刺激效应(P0.05),并且无论冻结与否,温带森林土壤微生物呼吸和微生物碳对外源Glu的响应均与植被类型、土壤湿度、外源氮供给及其形态存在显著的相关性。     

Impacts of an anomalously warm year on soil CO2 efflux in experimentally manipulated tallgrass prairie ecosystems

Modeling analyses suggest that an increase in growth rate of atmospheric CO₂ concentrations during an anomalously warm year may be caused by a decrease in net ecosystem production (NEP) in response to increased heterotrophic respiration (R_h). To test this hypothesis, 12 intact soil monoliths were excavated from a tallgrass prairie site near Purcell, Oklahoma, USA and divided among four large dynamic flux chambers (Ecologically Controlled Enclosed Lysimeter Laboratories (EcoCELLs)). During the first year, all four EcoCELLs were subjected to Oklahoma air temperatures. During the second year, air temperature in two EcoCELLs was increased by 4°C throughout the year to simulate anomalously warm conditions. This paper reports on the effect of warming on soil CO₂ efflux, representing the sum of autotrophic respiration (R_a) and R_h. During the pretreatment year, weekly average soil CO₂ efflux was similar in all EcoCELLs. During the late spring, summer and early fall of the treatment year, however, soil CO₂ efflux was significantly lower in the warmed EcoCELLs. In general, soil CO₂ efflux was correlated with soil temperature and to a lesser extent with moisture. A combined temperature and moisture regression explained 64% of the observed variation in soil CO₂ efflux. Soil CO₂ efflux correlated well with a net primary production (NPP) weighted greenness index derived from digital photographs. Although separate relationships for control and warmed EcoCELLs showed better correlations, one single relationship explained close to 70% of the variation in soil CO₂ efflux across treatments and years. A strong correlation between soil CO₂ efflux and canopy development and the lack of initial response to warming indicate that soil CO₂ efflux is dominated by R_a. This study showed that a decrease in soil CO₂ efflux in response to a warm year was most likely dominated by a decrease in R_a instead of an increase in R_h.     

Quantitative Isolation of Microbial DNA from Different Types of Soils of Natural and Agricultural Ecosystems

A novel procedure was developed for direct quantitative isolation of microbial DNA from soil. This technique was used to evaluate microbial DNA pools in soils of contrasting types (chernozems and brown forest soils) under different anthropogenic loads. A strong correlation was found between microbial biomass and DNA contents in soils of different types (R ²= 0.799). The ratio of soil CO₂ emission rate to the amount of extractable DNA in the soil was shown to reflect the physiological state of the soil microbial community; this ratio can be used as an ecophysiological parameter similarly to the metabolic quotient qCO₂.     

Effects of an induced drought on soil carbon dioxide (CO2) efflux and soil CO2 production in an Eastern Amazonian rainforest, Brazil

In the next few decades, climate of the Amazon basin is expected to change, as a result of deforestation and rising temperatures, which may lead to feedback mechanisms in carbon (C) cycling that are presently unknown. Here, we report how a throughfall exclusion (TFE) experiment affected soil carbon dioxide (CO₂) production in a deeply weathered sandy Oxisol of Caxiuanã (Eastern Amazon). Over the course of 2 years, we measured soil CO₂ efflux and soil CO₂ concentrations, soil temperature and moisture in pits down to 3 m depth. Over a period of 2 years, TFE reduced on average soil CO₂ efflux from 4.3±0.1 μmol CO₂ m⁻² s⁻¹ (control) to 3.2±0.1 μmol CO₂ m⁻² s⁻¹ (TFE). The contribution of the subsoil (below 0.5 m depth) to the total soil CO₂ production was higher in the TFE plot (28%) compared with the control plot (17%), and it did not differ between years. We distinguished three phases of drying after the TFE was started. The first phase was characterized by a translocation of water uptake (and accompanying root activity) to deeper layers and not enough water stress to affect microbial activity and/or total root respiration. During the second phase a reduction in total soil CO₂ efflux in the TFE plot was related to a reduction of soil and litter decomposers activity. The third phase of drying, characterized by a continuing decrease in soil CO₂ production was dominated by a water stress‐induced decrease in total root respiration. Our results contrast to results of a drought experiment on clay Oxisols, which may be related to differences in soil water retention characteristics and depth of rooting zone. These results show that large differences exist in drought sensitivity among Amazonian forest ecosystems, which primarily seem to be affected by the combined effects of texture (affecting water holding capacity) and depth of rooting zone.     

Carbon balance and water use efficiency of frequently cut Lolium perenne L. swards at elevated carbon dioxide

The impact of doubled atmospheric [CO₂] on the carbon balance of regularly cut Lolium perenne L. swards was studied for two years under semi-field conditions in the Wageningen Rhizolab. CO₂ and H₂O vapour exchange rates of the swards were measured continuously for two years in transparent enclosures. The light utilization efficiencies of the swards ranged between 1.5 g CO₂ MJ^–1 global radiation (high light, ambient [CO₂]) and 2.8 g CO₂ MJ^–1 (low light, doubled [CO₂]). The above-ground net primary productivity (NPP) in the enclosures was greater by 29% in 1994 and 43% in 1995 in the doubled [CO₂] treatments, but only 20% and 25% more carbon was recovered in the periodical cuts. Thus, NPP increased significantly more than did the harvested above-ground biomass. The positive [CO₂] effect on net carbon assimilation is therefore associated with a preferential allocation of extra carbon to the roots and soil. In addition to higher canopy photosynthesis and leaf elongation rates, a small part of the positive [CO₂] effects on NPP could be attributed to a decrease of the specific respiration of the shoots. On a canopy basis however, respiration was equal or slightly higher at doubled [CO₂] due to the higher amount of standing biomass. Comparison of NPP and carbon recovered in different harvests showed that allocation to roots and soil was highest in spring, it was low in early summer and increased again in late summer and autumn. The total gross amount of carbon partitioned to the roots and soil during the two year period was 57% more at doubled [CO₂]. The total amount of carbon that was sequestered in the soil after subtraction of the respiratory losses was 458 g m^–2 and 779 g m^–2 in the ambient and doubled [CO₂] treatments, respectively. The average water use efficiency (WUE) of the swards was increased by a factor 1.5 at doubled [CO₂]. Both WUE and its positive interaction with [CO₂] varied between years and were positively correlated with global irradiance. At doubled [CO₂], the higher WUE was fully compensated for by a higher leaf area index. Therefore, total transpiration on a canopy basis was equal for the ambient and the doubled [CO₂] concentrations in both years.     

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

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

Widespread foliage δ15N depletion under elevated CO2: inferences for the nitrogen cycle

Leaf ¹⁵N signature is a powerful tool that can provide an integrated assessment of the nitrogen (N) cycle and whether it is influenced by rising atmospheric CO₂ concentration. We tested the hypothesis that elevated CO₂ significantly changes foliage δ¹⁵N in a wide range of plant species and ecosystem types. This objective was achieved by determining the δ¹⁵N of foliage of 27 field‐grown plant species from six free‐air CO₂ enrichment (FACE) experiments representing desert, temperate forest, Mediterranean‐type, grassland prairie, and agricultural ecosystems. We found that within species, the δ¹⁵N of foliage produced under elevated CO₂ was significantly lower (P<0.038) compared with that of foliage grown under ambient conditions. Further analysis of foliage δ¹⁵N by life form and growth habit revealed that the CO₂ effect was consistent across all functional groups tested. The examination of two chaparral shrubs grown for 6 years under a wide range of CO₂ concentrations (25–75 Pa) also showed a significant and negative correlation between growth CO₂ and leaf δ¹⁵N. In a select number of species, we measured bulk soil δ¹⁵N at a depth of 10 cm, and found that the observed depletion of foliage δ¹⁵N in response to elevated CO₂ was unrelated to changes in the soil δ¹⁵N. While the data suggest a strong influence of elevated CO₂ on the N cycle in diverse ecosystems, the exact site(s) at which elevated CO₂ alters fractionating processes of the N cycle remains unclear. We cannot rule out the fact that the pattern of foliage δ¹⁵N responses to elevated CO₂ reported here resulted from a general drop in δ¹⁵N of the source N, caused by soil‐driven processes. There is a stronger possibility, however, that the general depletion of foliage δ¹⁵N under high CO₂ may have resulted from changes in the fractionating processes within the plant/mycorrhizal system.     

Does the lichen mat alleviate the effects of wet deposited nickel on soil microorganisms and Scots pine (Pinus sylvestris L.) seedlings?

A field experiment was conducted in a dry heath forest dominated by Scots pine (Pinus sylvestris L.) and a mat-forming lichen (Cladina stellaris (Opiz) Brodo) to assess the effect of wet-deposited nickel (Ni) on pine seedlings and soil microorganisms, and to explore whether an intact lichen mat could act as a buffer against heavy metal deposits. Pine seedlings were planted in quadrats covered by a natural lichen layer and in quadrats from which the lichen layer had been completely removed. The quadrats were exposed to four levels of Ni deposition: 0 (i.e., distilled water), 10, 100 and 1000 mg m^–2 year^–1 in two growing seasons. Increasing Ni deposition led to an increase in the Ni content of the needles, roots and the soil organic layer. The lichen mat reduced Ni flow to the organic soil layer, but had no significant, reducing effect on needle or root Ni concentration. The most severe Ni treatment had detrimental effects on seedling growth and increased peroxidase activity in the previous years needles. Removal of the lichen layer did not increase susceptibility of seedlings to Ni. Values of maximal carbon use efficiency (Max) and metabolic quotient (qCO₂) of the soil microorganisms indicated protective value of the lichen mat to soil microorganisms at the highest Ni treatment. Skimming per se decreased basal respiration, qCO₂ and concentrations of potassium in the soil and also increased the lag period of the microorganisms as a response to in situ substrate addition.     

Characterization of the non-photochemical quenching of chlorophyll fluorescence that occurs during the active accumulation of inorganic carbon in the cyanobacterium Synechococcus PCC 7942

Previous work has shown that the maximum fluorescence yield from PS 2 of Synechococcus PCC 7942 occurs when the cells are at the CO₂ compensation point. The addition of inorganic carbon (C_i), as CO₂ or HCO₃ ^–, causes a lowering of the fluorescence yield due to both photochemical (q_p) and non-photochemical (q_N) quenching. In this paper, we characterize the q_N that is induced by C_i addition to cells grown at high light intensities (500 mol photons m^–2 s^–1). The C_i-induced q_N was considerably greater in these cells than in cells grown at low light intensities (50 mol photons m^–2 s^–1), when assayed at a white light (WL) intensity of 250 mol photons m^–2 s^–1. In high-light grown cells we measured q_N values as high as 70%, while in low-light grown cells the q_N was about 16%. The q_N was relieved when cells regained the CO₂ compensation point, when cells were illuminated by supplemental far-red light (FRL) absorbed mainly by PS 1, or when cells were illuminated with increased WL intensities. These characteristics indicate that the q_N was not a form of energy quenching (q_E). Supplemental FRL illumination caused significant enhancement of photosynthetic O₂ evolution that could be correlated with the changes in q_p and q_N. The increases in q_p induced by C_i addition represent increases in the effective quantum yield of PS 2 due to increased levels of oxidized Q_A. The increase in q_N induced by C_i represents a decrease in PS 2 activity related to decreases in the potential quantum yield. The lack of diagnostic changes in the 77 K fluorescence emission spectrum argue against q_N being related to classical state transitions, in which the decrease in potential quantum yield of PS 2 is due either to a decrease in absorption cross-section or by increased spill-over of excitation energy to PS 1. Both the C_i-induced q_p (t _0.5<0.5 s) and q_N (t _0.51.6 s) were rapidly relieved by the addition of DCMU. The two time constants give further support for two separate quenching mechanisms. We have thus characterized a novel form of q_N in cyanobacteria, not related to state transitions or energy quenching, which is induced by the addition of C_i to cells at the CO₂-compensation point.Abbreviations BTP- 1,3-bis[tris(hydroxymethyl)-methylaminopropane] - Chl- chlorophyll - C_i- inorganic carbon (CO₂+HCO₃ ^–+CO₃ ^2–) - DCMU- 3-(3,4-dichlorophenyl)-, 1-dimethylurea) - F- chlorophyll fluorescence measured at any time in the absence of a saturating flash - F_o- chlorophyll fluorescence with only the weak modulated measuring beam on - F_M'- chlorophyll fluorescence during a saturating flash - F_M- maximum chlorophyll fluorescence, measured in the presence of WL and FRL at the CO₂-compensation point or in the presence of DCMU - F_V- variable fluorescence (= F_M'–F₀) - FRL- supplemental illumination with far red light - MB- modulated measuring beam of the PAM fluorometer - MV- methyl viologen - PAM- pulse amplitude modulation - PFD- incident photon flux density - PS 1, 2- Photosystems 1 and 2 - Q_A- primary electron-accepting plastoquinione of PS 2 - q_N- non-photochemical quenching of chlorophyll fluorescence - q_p- photochemical quenching of chlorophyll fluorescence; rubisco-ribulose bisphosphate carboxylase/oxygenase - SF- saturating flash (600 ms duration) - WL- white light illumination     

亚热带5个生态系统土壤微生物学性质比较

为了解亚热带生态系统土壤微生物学特性,对疏草荒地(Barren grassland)、纯杉木林(Cunninghamia lanceolata forest)、纯樟树林(Cinnamomum camphora forest)、杉木樟树混交林(Cunninghamia lanceolata-Cinnamomum camphora mixed forest)和自然恢复地(Nature recovery)等5种生态系统土壤微生物学性质进行了分析。结果表明,在0~20、20~40和40~60 cm土层,纯樟树林、杉木樟树混交林和自然恢复地的土壤微生物生物量碳氮磷和土壤基础呼吸显著高于疏草荒地和纯杉木林(P0.05),而他们的代谢熵低于疏草荒地和纯杉木林(P0.05);随着土层深度的增加,不同生态系统的土壤微生物生物量碳氮磷和土壤基础呼吸显著减小(P0.05),代谢熵显著增大(P0.05)。土壤容重与土壤微生物生物量碳氮磷和土壤基础呼吸呈显著负相关(P0.05),与代谢熵呈显著正相关(P0.05);土壤微生物生物量碳氮磷和土壤基础呼吸均与土壤总孔隙度、土壤水稳性大团聚体、土壤有机碳、全氮、有效氮、有效磷呈极显著正相关(P0.01),代谢熵与土壤总孔隙度、土壤水稳性大团聚体、土壤有机碳、全氮、有效氮和有效磷呈显著负相关(P0.05)。因此,自然恢复地、纯樟树林和杉木樟树混交林对土壤质量的改善有明显促进作用,而疏草荒地和纯杉木林对土壤改良效果不明显。     

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.     

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.     

Fine root dynamics in a loblolly pine forest are influenced by free-air-CO2-enrichment: a six-year-minirhizotron study

Efforts to characterize carbon (C) cycling among atmosphere, forest canopy, and soil C pools are hindered by poorly quantified fine root dynamics. We characterized the influence of free‐air‐CO₂‐enrichment (ambient +200 ppm) on fine roots for a period of 6 years (Autumn 1998 through Autumn 2004) in an 18‐year‐old loblolly pine (Pinus taeda) plantation near Durham, NC, USA using minirhizotrons. Root production and mortality were synchronous processes that peaked most years during spring and early summer. Seasonality of fine root production and mortality was not influenced by atmospheric CO₂ availability. Averaged over all 6 years of the study, CO₂ enrichment increased average fine root standing crop (+23%), annual root length production (+25%), and annual root length mortality (+36%). Larger increase in mortality compared with production with CO₂ enrichment is explained by shorter average fine root lifespans in elevated plots (500 days) compared with controls (574 days). The effects of CO₂‐enrichment on fine root proliferation tended to shift from shallow (0–15 cm) to deeper soil depths (15–30) with increasing duration of the study. Diameters of fine roots were initially increased by CO₂‐enrichment but this effect diminished over time. Averaged over 6 years, annual fine root NPP was estimated to be 163 g dw m^?2 yr^?1 in CO₂‐enriched plots and 130 g dw m^?2 yr^?1 in control plots (P= 0.13) corresponding to an average annual additional input of fine root biomass to soil of 33 g m^?2 yr^?1 in CO₂‐enriched plots. A lack of consistent CO₂× year effects suggest that the positive effects of CO₂ enrichment on fine root growth persisted 6 years following minirhizotron tube installation (8 years following initiation of the CO₂ fumigation). Although CO₂‐enrichment contributed to extra flow of C into soil in this experiment, the magnitude of the effect was small suggesting only modest potential for fine root processes to directly contribute to soil C storage in south‐eastern pine forests.     

Elevated atmospheric CO2 effects on biomass production and soil carbon in conventional and conservation cropping systems

Increasing atmospheric CO₂ concentration has led to concerns about potential effects on production agriculture as well as agriculture's role in sequestering C. In the fall of 1997, a study was initiated to compare the response of two crop management systems (conventional and conservation) to elevated CO₂. The study used a split‐plot design replicated three times with two management systems as main plots and two CO₂ levels (ambient=375 μL L^?1 and elevated CO₂=683 μL L^?1) as split‐plots using open‐top chambers on a Decatur silt loam (clayey, kaolinitic, thermic Rhodic Paleudults). The conventional system was a grain sorghum (Sorghum bicolor (L.) Moench.) and soybean (Glycine max (L.) Merr.) rotation with winter fallow and spring tillage practices. In the conservation system, sorghum and soybean were rotated and three cover crops were used (crimson clover (Trifolium incarnatum L.), sunn hemp (Crotalaria juncea L.), and wheat (Triticum aestivum L.)) under no‐tillage practices. The effect of management on soil C and biomass responses over two cropping cycles (4 years) were evaluated. In the conservation system, cover crop residue (clover, sunn hemp, and wheat) was increased by elevated CO₂, but CO₂ effects on weed residue were variable in the conventional system. Elevated CO₂ had a greater effect on increasing soybean residue as compared with sorghum, and grain yield increases were greater for soybean followed by wheat and sorghum. Differences in sorghum and soybean residue production within the different management systems were small and variable. Cumulative residue inputs were increased by elevated CO₂ and conservation management. Greater inputs resulted in a substantial increase in soil C concentration at the 0–5 cm depth increment in the conservation system under CO₂‐enriched conditions. Smaller shifts in soil C were noted at greater depths (5–10 and 15–30 cm) because of management or CO₂ level. Results suggest that with conservation management in an elevated CO₂ environment, greater residue amounts could increase soil C storage as well as increase ground cover.     

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.     

Effects of elevated atmospheric CO2 on fine root production and activity in an intact temperate forest ecosystem

We investigated the effects of elevated atmospheric CO₂ concentrations (ambient + 200 ppm) on fine root production and soil carbon dynamics in a loblolly pine (Pinus taeda) forest subject to free‐air CO₂ enrichment (FACE) near Durham, NC (USA). Live fine root mass (LFR) showed less seasonal variation than dead fine root mass (DFR), which was correlated with seasonal changes in soil moisture and soil temperature. LFR mass increased significantly (by 86%) in the elevated CO₂ treatment, with an increment of 37 g(dry weight) m^?2 above the control plots after two years of CO₂ fumigation. There was no long‐term increment in DFR associated with elevated CO₂, but significant seasonal accumulations of DFR mass occurred during the summer of the second year of fumigation. Overall, root net primary production (RNPP) was not significantly different, but annual carbon inputs were 21.7 gC m^?2 y^?1 (68%) higher in the elevated CO₂ treatment compared to controls. Specific root respiration was not altered by the CO₂ treatment during most of the year; however, it was significantly higher by 21% and 13% in September 1997 and May 1998, respectively, in elevated CO₂. We did not find statistically significant differences in the C/N ratio of the root tissue, root decomposition or phosphatase activity in soil and roots associated with the treatment. Our data show that the early response of a loblolly pine forest ecosystem subject to CO₂ enrichment is an increase in its fine root population and a trend towards higher total RNPP after two years of CO₂ fumigation.     

The effects of free-air CO2 enrichment and soil water availability on spatial and seasonal patterns of wheat root growth

Spring wheat [ Triticum aestivum (L). cv. Yecora Rojo] was grown from December 1992 to May 1993 under two atmospheric CO₂ concentrations, 550 μmol mol^–1 for high-CO₂ plots, and 370 μmol mol^–1 for control plots, using a Free-Air CO₂ Enrichment (FACE) apparatus. In addition to the two levels of atmospheric CO₂, there were ample and limiting levels of water supply through a subsurface trip irrigation system in a strip, split-plot design. In order to examine the temporal and spatial root distribution, root cores were extracted at six growth stages during the season at in-row and inter-row positions using a soil core device (86 mm ID, 1.0 m length). Such information would help determine whether and to what extent root morphology is changed by alteration of two important factors, atmospheric CO₂ and soil water, in this agricultural ecosystem. Wheat root growth increased under elevated CO₂ conditions during all observed developmental stages. A maximum of 37% increase in total root dry mass in the FACE vs. Control plots was observed during the period of stem elongation. Greater root growth rates were calculated due to CO₂ enhancement until anthesis. During the early vegetative growth, root dry mass of the inter-row space was significantly higher for FACE than for Control treatments suggesting that elevated CO₂ promoted the production of first-order lateral roots per main axis. Then, during the reproductive period of growth, more branching of lateral roots in the FACE treatment occurred due to water stress. Significant higher root dry mass was measured in the inter-row space of the FACE plots where soil water supply was limiting. These sequential responses in root growth and morphology to elevated CO₂ and reduced soil water supports the hypothesis that plants grown in a high-CO₂ environment may better compensate soil-water-stress conditions.     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

Subsurface CO<Subscript>2</Subscript> Dynamics in Temperate Beech and Spruce Forest Stands

Rates of soil respiration (CO₂ effluxes), subsurface pore gas CO₂/O₂ concentrations, soil temperature and soil water content were measured for 15 months in two temperate and contrasting Danish forest ecosystems: beech (Fagus sylvatica L.) and Norway spruce (Picea abies [L.] Karst.). Soil CO₂ effluxes showed a distinct seasonal trend in the range of 0.48–3.3 μmol CO₂ m⁻² s⁻¹ for beech and 0.50–2.92 μmol CO₂ m⁻² s⁻¹ for spruce and were well-correlated with near-surface soil temperatures. The soil organic C-stock (upper 1 m including the O-horizon) was higher in the spruce stand (184±23 Mg C ha⁻¹) compared to the beech stand (93±19 Mg C ha⁻¹) and resulted in a faster turnover time as calculated by mass/flux in soil beneath the beech stand (28 years) compared to spruce stand (60 years). Observed soil CO₂ concentrations and effluxes were simulated using a Fickian diffusion-reaction model based on vertical CO₂ production rates and soil diffusivity. Temporal trends were simulated on the basis of observed trends in the distribution of soil water, temperature, and live roots as well as temperature and water content sensitivity functions. These functions were established based on controlled laboratory incubation experiments. The model was successfully validated against observed soil CO₂ effluxes and concentrations and revealed that temporal trends generally could be linked to variations in subsurface CO₂ production rates and diffusion over time and with depths. However, periods with exceptionally high CO₂ effluxes (> 20 μmol CO₂ m⁻² s⁻¹) were noted in March 2000 in relation to drying after heavy rain and after the removal of snow from collars. Both cases were considered non-steady state and could not be simulated.