Dynamics of soil CO2 efflux under varying atmospheric CO2 concentrations reveal dominance of slow processes

We evaluated the effect on soil CO₂ efflux (F_CO2) of sudden changes in photosynthetic rates by altering CO₂ concentration in plots subjected to +200 ppmv for 15 years. Five‐day intervals of exposure to elevated CO₂ (eCO₂) ranging 1.0–1.8 times ambient did not affect F_CO2. F_CO2 did not decrease until 4 months after termination of the long‐term eCO₂ treatment, longer than the 10 days observed for decrease of F_CO2 after experimental blocking of C flow to belowground, but shorter than the ~13 months it took for increase of F_CO2 following the initiation of eCO₂. The reduction of F_CO2 upon termination of enrichment (~35%) cannot be explained by the reduction of leaf area (~15%) and associated carbohydrate production and allocation, suggesting a disproportionate contraction of the belowground ecosystem components; this was consistent with the reductions in base respiration and F_CO2‐temperature sensitivity. These asymmetric responses pose a tractable challenge to process‐based models attempting to isolate the effect of individual processes on F_CO2.     

The effects of elevated CO2 on symbiotic N2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems

The response of plants to elevated CO₂ is dependent on the availability of nutrients, especially nitrogen. It is generally accepted that an increase in the atmospheric CO₂ concentration increases the C:N ratio of plant residues and exudates. This promotes temporary N-immobilization which might, in turn, reduce the availability of soil nitrogen. In addition, both a CO₂ stimulated increase in plant growth (thus requiring more nitrogen) and an increased N demand for the decomposition of soil residues with a large C:N will result under elevated CO₂ in a larger N-sink of the whole grassland ecosystem. One way to maintain the balance between the C and N cycles in elevated CO₂ would be to increase N-import to the grassland ecosystem through symbiotic N₂ fixation. Whether this might happen in the context of temperate ecosystems is discussed, by assessing the following hypothesis: i) symbiotic N₂ fixation in legumes will be enhanced under elevated CO₂, ii) this enhancement of N₂ fixation will result in a larger N-input to the grassland ecosystem, and iii) a larger N-input will allow the sequestration of additional carbon, either above or below-ground, into the ecosystem. Data from long-term experiments with model grassland ecosystems, consisting of monocultures or mixtures of perennial ryegrass and white clover, grown under elevated CO₂ under free-air or field-like conditions, supports the first two hypothesis, since: i) both the percentage and the amount of fixed N increases in white clover grown under elevated CO₂, ii) the contribution of fixed N to the nitrogen nutrition of the mixed grass also increases in elevated CO₂. Concerning the third hypothesis, an increased nitrogen input to the grassland ecosystem from N₂ fixation usually promotes shoot growth (above-ground C storage) in elevated CO₂. However, the consequences of this larger N input under elevated CO₂ on the below-ground carbon fluxes are not fully understood. On one hand, the positive effect of elevated CO₂ on the quantity of plant residues might be overwhelming and lead to an increased long-term below-ground C storage; on the other hand, the enhancement of the decomposition process by the N-rich legume material might favour carbon turn-over and, hence, limit the storage of below-ground carbon.     

Ecosystem constraints to symbiotic nitrogen fixers: a simple model and its implications

The widespread occurrence of N limitation to net primary production (NPP) and other ecosystem processes, despite the ubiquitous occurrence of N-fixing symbioses, remains a significant puzzle in terrestrial ecology. We describe a simple simulation model for an ecosystem containing a generic nonfixer and a symbiotic N fixer, based on: (1) a higher cost for N acquisition by N fixers than nonfixers; (2) growth of fixers and fixation of N only when low N availability limits the growth of nonfixers, and other resources are available; and (3) losses of fixed N from the system only when the quantity of available N exceeds plant and microbial demands. Despite the disadvantages faced by the N fixer under these conditions, N fixation and loss adjust N availability close to the availability of other resources, and biomass and NPP in this simple model can be substantially but only transiently N limited. We then modify the model by adding: (1) losses of N in forms other than excess available N (e.g., dissolved organic N, trace gases produced by nitrification); and (2) constraints to the growth and activity of N fixers imposed by differential effects of shading, P limitation, and grazing. The combination of these processes is sufficient to describe an open system, with input from both precipitation and N fixation, that is nevertheless strongly N-limited at equilibrium. This model is useful for exploring causes and consequences of constraints to N fixation, and hence of N limitation, and we believe it will also be useful for evaluating how N fixation and limitation interact with elevated CO₂ and other components of global enviromental change.     

Constraints to nitrogen acquisition of terrestrial plants under elevated CO2

A key part of the uncertainty in terrestrial feedbacks on climate change is related to how and to what extent nitrogen (N) availability constrains the stimulation of terrestrial productivity by elevated CO₂ (eCO₂), and whether or not this constraint will become stronger over time. We explored the ecosystem‐scale relationship between responses of plant productivity and N acquisition to eCO₂ in free‐air CO₂ enrichment (FACE) experiments in grassland, cropland and forest ecosystems and found that: (i) in all three ecosystem types, this relationship was positive, linear and strong (r² = 0.68), but exhibited a negative intercept such that plant N acquisition was decreased by 10% when eCO₂ caused neutral or modest changes in productivity. As the ecosystems were markedly N limited, plants with minimal productivity responses to eCO₂ likely acquired less N than ambient CO₂‐grown counterparts because access was decreased, and not because demand was lower. (ii) Plant N concentration was lower under eCO₂, and this decrease was independent of the presence or magnitude of eCO₂‐induced productivity enhancement, refuting the long‐held hypothesis that this effect results from growth dilution. (iii) Effects of eCO₂ on productivity and N acquisition did not diminish over time, while the typical eCO₂‐induced decrease in plant N concentration did. Our results suggest that, at the decennial timescale covered by FACE studies, N limitation of eCO₂‐induced terrestrial productivity enhancement is associated with negative effects of eCO₂ on plant N acquisition rather than with growth dilution of plant N or processes leading to progressive N limitation.     

Decomposition and nitrogen release from leaves of three hardwood species grown under elevated O3 and/or CO2

Elevated concentrations of O₃ and CO₂ have both been shown to affect structure, nutrient status, and deposition of secondary metabolites in leaves of forest trees. While such studies have produced robust models of the effects of such air pollutants on tree ecophysiology and growth, few have considered the potential for broader, ecosystem-level effects after these chemically and structurally altered leaves fall as leaf litter and decay. To determine the effects of elevated O₃ and/or CO₂ on the subsequent decomposition and nutrient release from the leaves grown in such altered atmospheres, we grew seedlings of three widespread North American forest trees, black cherry (Prunus serotina) (BC), sugar maple (Acer saccharum) (SM), and yellow-poplar (Liriodendron tulipifera) (YP) for two growing seasons in charcoal-filtered air (CF-air=approximately 25% ambient O₃), ambient O₃ (1X) or twice-ambient O₃ (2X) in outdoor open-top chambers. We then assayed the loss of mass and N from the litter derived from those seedlings through one year litterbag incubations in the forest floor of a neighboring forest stand. Mass loss followed linear functions and was not affected by the O₃ regime in which the leaves were grown. Instantaneous decay rates (i.e. k values) averaged SM:–0.707 y^-1, BC:–0.613 y^-1, and YP:–0.859 y^-1. N loss from ambient (1X) O₃-grown SM leaves was significantly greater than from CF-air leaves: N loss from BC leaves did not differ among treatments. Significantly less N was released from CF-air-grown YP leaves than from 1X or 2X O₃-treated leaves. YP leaves from plants grown in pots at 2X O₃ and 350 ppm supplemental CO₂ in indoor pollutant fumigation chambers (CSTRs or Continuously Stirred Tank Reactors) loss 40% as much mass and 27% as much N over one year as did leaves from YP grown in CF-air or 2X O₃. Thus, for leaves from plants grown in pots in controlled environment fumigation chambers, the concentrations of both O₃ and CO₂ can affect N release from litter incubated in the field whereas mass loss rate was affected only by CO₂. Because both mass loss and N release from leaves grown at elevated CO₂ were reduced significantly (at least for yellow-poplar), forests exposed to elevated CO₂ may have significantly reduced N turnover rates, thereby resulting in increased N limitation of tree growth, especially in forests which are already N-limited.     

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Carbon and nitrogen allocation in Lolium perenne in response to elevated atmospheric CO2 with emphasis on soil carbon dynamics

The effect of elevated CO2 on the carbon and nitrogen distribution within perennial ryegrass (L. perenne L.) and its influence on belowground processes were investigated. Plants were homogeneously 14C-labelled in two ESPAS growth chambers in a continuous 14C-CO2 atmosphere of 350 and 700 L L-1 CO2 and at two soil nitrogen regimes, in order to follow the carbon flow through all plant and soil compartments.After 79 days, elevated CO2 increased the total carbon uptake by 41 and 21% at low (LN) and high nitrogen (HN) fertilisation, respectively. Shoot growth remained unaffected, whereas CO2 enrichment stimulated root growth by 46% and the root/soil respiration by 111%, irrespective of the nitrogen concentration. The total 14C-soil content increased by 101 and 28% at LN and HN, respectively. The decomposition of the native soil organic matter was not affected either by CO2 or by the nitrogen treatment.Elevated CO2 did not change the total nitrogen uptake of the plant either at LN or at HN. Both at LN and HN elevated CO2 significantly increased the total amount of nitrogen taken up by the roots and decreased the absolute and relative amounts translocated to the shoots.The amount of soil nitrogen immobilised by micro-organisms and the size of the soil microbial biomass were not affected by elevated CO2, whereas both were significantly increased at the higher soil N content.Most striking was the 88% increase in net carbon input into the soil expressed as: 14C-roots plus total 14C-soil content minus the 12C-carbon released by decomposition of native soil organic matter. The net carbon input into the soil at ambient CO2 corresponded with 841 and 1662 kg ha-1 at LN and HN, respectively. Elevated CO2 increased these amounts with an extra carbon input of 950 and 1056 kg ha-1. Combined with a reduced decomposition rate of plant material grown at elevated CO2 this will probably lead to carbon storage in grassland soils resulting in a negative feed back on the increasing CO2 concentration of the atmosphere.     

大气CO2浓度升高对不同施氮土壤酶活性的影响

利用中国唯一的无锡FACE（Free-air CO2 enrichment,开放式空气CO2浓度升高）平台,研究了大气CO2浓度升高对土壤β-葡糖苷酶、转化酶、脲酶、酸性磷酸酶、-氨基葡糖苷酶的影响。研究发现,不同氮肥处理下大气CO2浓度升高对某些土壤酶活性的影响不同。在低氮施肥处理中,大气CO2浓度升高显著降低-葡糖苷酶活性,但是在高氮施肥处理下,大气CO2浓度升高显著增加β-葡糖苷酶活性。在低氮和常氮施肥处理中大气CO2浓度升高显著增加了土壤脲酶活性,但在高氮水平下影响不显著。在低氮、常氮施肥处理中,大气CO2浓度升高对土壤酸性磷酸酶活性没有影响,而在高氮施肥处理中显著增强了土壤中磷酸酶活性。大气CO2浓度升高对土壤转化酶活性和-氨基葡糖苷酶的活性有增加趋势,但影响不显著。研究还发现,在不同的CO2浓度下,土壤酶活性对不同氮肥处理的响应也不同。在正常CO2浓度下,土壤中β-葡糖苷酶活性随着氮肥施用量的增加而降低,而在大气CO2浓度升高条件下,却随着氮肥施用量的增加而增加。在大气CO2浓度升高条件下,高氮施肥显著增加了转化酶和酸性磷酸酶活性,而在正常CO2浓度下,影响不显著。在大气CO2浓度升高条件下,氮肥处理对脲酶活性的影响不大,但在正常CO2浓度下,脲酶活性随着氮肥施用量的增加而增加。氮肥对β-氨基葡糖苷酶活性的影响不明显。     

Predicting responses of photosynthesis and root fraction to elevated [CO2]a: interactions among carbon,nitrogen, and growth*

At elevated atmospheric CO₂ concentrations ([CO₂]_a), photosynthetic capacity (A_max) and root fraction (η_R, the ratio of root to plant dry mass) increased in some studies and decreased in others. Here, we have explored possible causes of this, focusing on the relative magnitudes of the effects of elevated [CO₂]_a on specific leaf (n_m) and plant (n_p) nitrogen concentrations, leaf mass per unit area (h), and plant nitrogen productivity (α). In our survey of 39 studies with 35 species, we found that elevated [CO₂]_a led to decreased n_m and n_p in all the studies and to increased h and α in most of the studies. The magnitudes of these changes varied with species and with experimental conditions. Based on a model that integrated [CO₂]_a-induced changes in leaf nitrogen into a biochemically based model of leaf photosynthesis, we predicted that, to a first approximation, photosynthesis will be upregulated (A_max will increase) when growth at increased [CO₂]_a leads to increases in h that are larger than decreases in n_m. Photosynthesis will be downregulated (A_max will decrease) when increases in h are smaller than decreases in n_m. The model suggests that photosynthetic capacity increases at elevated [CO₂]_a only when additional leaf mesophyll more than compensates the effects of nitrogen dilution. We considered two kinds of regulatory paradigms that could lead to varying responses of η_R to elevated [CO₂]_a, and compared the predictions of each with the data. A simple static model based on the functional balance concept predicts that η_R should increase when neither n_p nor h is very responsive to elevated [CO₂]_a. The quantitative and qualitative agreement of the predictions with data from the literature, however, is poor. A model that predicts η_R from the relative sensitivities of photosynthesis and relative growth rate to elevated [CO₂]_a corresponds much more closely to the observations. In general, root fraction increases if the response of photosynthesis to [CO₂]_a is greater than that of relative growth rate.     

Nitrogen application is required to realize wheat yield stimulation by elevated CO2 but will not remove the CO2‐induced reduction in grain protein concentration

Elevated CO₂ (eCO₂) generally promotes increased grain yield (GY) and decreased grain protein concentration (GPC), but the extent to which these effects depend on the magnitude of fertilization remains unclear. We collected data on the eCO₂ responses of GY, GPC and grain protein yield and their relationships with nitrogen (N) application rates across experimental data covering 11 field grown wheat (Triticum aestivum) cultivars studied in eight countries on four continents. The eCO₂‐induced stimulation of GY increased with N application rates up to ~200 kg/ha. At higher N application, stimulation of GY by eCO₂ stagnated or even declined. This was valid both when the yield stimulation was expressed as the total effect and using per ppm CO₂ scaling. GPC was decreased by on average 7% under eCO₂ and the magnitude of this effect did not depend on N application rate. The net effect of responses on GY and protein concentration was that eCO₂ typically increased and decreased grain protein yield at N application rates below and above ~100 kg/ha respectively. We conclude that a negative effect on wheat GPC seems inevitable under eCO₂ and that substantial N application rates may be required to sustain wheat protein yields in a world with rising CO₂.     

CO2浓度升高和施氮条件下小麦根际呼吸对土壤呼吸的贡献

依托FACE技术平台,采用稳定13C同位素技术,通过将小麦(C3作物)种植于长期单作玉米(C4作物)的土壤上,研究了大气CO2浓度升高和不同氮肥水平对土壤排放CO2的δ13C值及根际呼吸的影响.结果表明:种植小麦后土壤排放CO2的δ13C值随作物生长逐渐降低,CO2浓度升高200 μmol·mol-1显著降低了孕穗、抽穗期(施氮量为250 kg·hm-2,HN)与拔节、孕穗期(施氮量为150 kg·hm-2,LN)土壤排放CO2的δ13C值,显著提高了孕穗、抽穗期的根际呼吸比例.拔节至成熟期,根际呼吸占土壤呼吸的比例在高CO2浓度下为24％～48％ (HN)和21％ ～48％ (LN),在正常CO2浓度下为20％ ～36％ (HN)和19％～32％(LN).不同CO2浓度下土壤排放CO2的δ13C值和根际呼吸对氮肥增加的响应不同,CO2浓度与氮肥用量在拔节期对根际呼吸的交互效应显著.     

Lifetime and temporal occurrence of ectomycorrhizae on ponderosa pine (Pinus ponderosa Laws.) seedlings grown under varied atmospheric CO2 and nitrogen levels

Climate change (elevated atmospheric CO2, and altered air temperatures, precipitation amounts and seasonal patterns) may affect ecosystem processes by altering carbon allocation in plants, and carbon flux from plants to soil. Mycorrhizal fungi, as carbon sinks, are among the first soil biota to receive carbon from plants, and thereby influence carbon release from plants to soil. One step in this carbon release is via fine root and mycorrhizal turnover. It is necessary to know the lifetime and temporal occurrence of roots and mycorrhizae to determine the capacity of the soil ecosystem to sequester carbon assimilated aboveground. In this study, ponderosa pine (Pinus ponderosa Laws) seedlings were grown under three levels of atmospheric CO2 (ambient, 525 and 700 mol CO2 mol-1) and three levels of annual nitrogen additions (0,100 and 200 kg N ha-1) in open-top chambers. At a two-month frequency during 18 months, we observed ectomycorrhizal root tips observed using minirhizotron tubes and camera. The numbers of new mycorrhizal root tips, the numbers of tips that disappeared between two consecutive recording events, and the standing crop of tips at each event were determined. There were more mycorrhizal tips of all three types seen during the summer compared with other times of the year. When only the standing crop of mycorrhizal tips was considered, effects of the CO2 and N addition treatments on carbon allocation to mycorrhizal tips was weakly evident. However, when the three types of tips were considered collectively, tips numbers flux of carbon through mycorrhizae was greatest in the: (1) high CO2 treatment compared with the other CO2 treatments, and (2) intermediate N addition treatment compared with the other N addition treatments. A survival analysis on the entire 18 month cohort of tips was done to calculate the median lifetime of the mycorrhizal root tips. Average median lifetime of the mycorrhizal tips was 139 days and was not affected by nitrogen and CO2 treatments.     

Effects of nitrogen: phosphorus supply ratios on nitrogen fixation in agricultural and pastoral ecosystems

An analysis of data compiled from the literature confirms a strong inverse relationship between annual rates of nitrogen fixation and the soil nitrogen content in agricultural and pastoral ecosystems. However, this inverse relationship is strongly modified by the rate of application of phosphorus fertilizer, which strongly influences the activities of both symbiotic and non-symbiotic nitrogen fixing organisms. In the case of symbiotic legumes, the response of N-fixation to N and P is in part a result of changes in legume dominance within the plant community. These results, as well as supporting data presented from a review of experiments on nitrogen fixation in a variety of other terrestrial and aquatic ecosystems, provide important support for the hypothesis that phosphorus availability is a key regulator of nitrogen biogeochemistry. Published as Paper No. 9950, Journal Series, Nebraska Agricultural Research Division, University of Nebraska, Lincoln, NE, USA.     

Time-dependent responses of soil CO2 efflux components to elevated atmospheric [CO2] and temperature in experimental forest mesocosms

We previously used dual stable isotope techniques to partition soil CO₂ efflux into three source components (rhizosphere respiration, litter decomposition, and soil organic matter (SOM) oxidation) using experimental chambers planted with Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] seedlings. The components responded differently to elevated CO₂ (ambient + 200 mol mol^–1) and elevated temperature (ambient + 4 °C) treatments during the first year. Rhizosphere respiration increased most under elevated CO₂, and SOM oxidation increased most under elevated temperature. However, many studies show that plants and soil processes can respond to altered climates in a transient way. Herein, we extend our analysis to 2 years to evaluate the stability of the responses of the source components. Total soil CO₂ efflux increased significantly under elevated CO₂ and elevated temperature in both years (1994 and 1995), but the enhancement was much less in 1995. Rhizosphere respiration increased less under elevated temperature in 1995 compared with 1994. Litter decomposition also tended to increase comparatively less in 1995 under elevated CO₂, but was unresponsive to elevated temperature between years. In contrast, SOM oxidation was similar under elevated CO₂ in the 2 years. Less SOM oxidation occurred under elevated temperature in 1995 compared with 1994. Our results indicate that temporal variations can occur in CO₂ production by the sources. The variations likely involve responses to antecedent physical disruption of the soil and physiological processes.     

The photosynthesis – leaf nitrogen relationship at ambient and elevated atmospheric carbon dioxide: a meta-analysis

Estimation of leaf photosynthetic rate (A) from leaf nitrogen content (N) is both conceptually and numerically important in models of plant, ecosystem, and biosphere responses to global change. The relationship between A and N has been studied extensively at ambient CO₂ but much less at elevated CO₂. This study was designed to (i) assess whether the A–N relationship was more similar for species within than between community and vegetation types, and (ii) examine how growth at elevated CO₂ affects the A–N relationship. Data were obtained for 39 C3 species grown at ambient CO₂ and 10 C3 species grown at ambient and elevated CO₂. A regression model was applied to each species as well as to species pooled within different community and vegetation types. Cluster analysis of the regression coefficients indicated that species measured at ambient CO₂ did not separate into distinct groups matching community or vegetation type. Instead, most community and vegetation types shared the same general parameter space for regression coefficients. Growth at elevated CO₂ increased photosynthetic nitrogen use efficiency for pines and deciduous trees. When species were pooled by vegetation type, the A–N relationship for deciduous trees expressed on a leaf-mass basis was not altered by elevated CO₂, while the intercept increased for pines. When regression coefficients were averaged to give mean responses for different vegetation types, elevated CO₂ increased the intercept and the slope for deciduous trees but increased only the intercept for pines. There were no statistical differences between the pines and deciduous trees for the effect of CO₂. Generalizations about the effect of elevated CO₂ on the A–N relationship, and differences between pines and deciduous trees will be enhanced as more data become available.     

Optimal nitrogen allocation controls tree responses to elevated CO2

Despite the abundance of experimental data, understanding of forest responses to elevated CO2 is limited. Here I show that a key to previously unexplained production and leaf area responses lies in the interplay between whole-plant nitrogen (N) allocation and leaf photosynthesis. A simple tree growth model, controlled by net growth maximization through optimization of leaf area index (LAI) and plant N, is used to analyse CO2 responses in both young, expanding and closed, steady-state canopies. The responses are sensitive to only two independent parameters, the photosynthetic capacity per leaf N (a) and the fine-root N:leaf N ratio. The model explains observed CO2 responses of photosynthesis, production and LAI in four forest free air CO2 enrichment (FACE) experiments. Insensitivity of LAI except at low LAI, increase in light-use efficiency, and photosynthetic down-regulation (as a result of reduced leaf N per area) at elevated CO2 are all explained through the combined effects on a and leaf quantum efficiency. The model bridges the gap between the understanding of leaf-level and plant-level responses and provides a transparent framework for interpreting and linking structural (LAI) and functional (net primary production (NPP):gross primary production (GPP) ratio, light-use efficiency, photosynthetic down-regulation) responses to elevated CO2.     

Biomass and compositional responses of ectomycorrhizal fungal hyphae to elevated CO2 and nitrogen fertilization

The extramatrical mycelia (EMM) of ectomycorrhizal fungi make up a large proportion of the microbial diversity and biomass in temperate forest soils. Thus, their response to elevated CO(2) can have large effects on plant nutrient acquisition and carbon movement through forests. Here, the effects of CO(2) and nitrogen (N) fertilization on EMM biomass and community structure in Pinus taeda forest plots were examined using sand-filled mesh bags buried in the field, the contents of which were analyzed by phospholipid fatty acid (PLFA) and DNA sequencing. A total of 2138 sequences comprising 295 taxa were recovered; most (83.5%) were from ectomycorrhizal fungal taxa. No biomass increase was detected in elevated CO(2) plots relative to control plots, but individual taxa responded to both CO(2) and N fertilization, four of the six most abundant taxa were less frequent in N-fertilized plots. Thelephoroid and athelioid taxa were both frequent and abundant as EMM, and thelephoroid richness was extremely high. Russula and Cortinariaceae taxa were less abundant and boletoid taxa were more abundant as EMM relative to ectomycorrhizas. The EMM community, sampled across seasons and years, was dynamic with a high degree of interspecific variation in response to CO(2) enrichment and N fertilization.     

Evaluation of symbiotic properties and nitrogen contribution of mucuna to maize grown in the derived savanna of West Africa

The severity and increase of the Imperata cylindrica constraint as a weed, the decline of the traditional fallow systems as a means of soil fertility management and the lack of inorganic fertilizer appear to have created opportunities for adoption of mucuna (Mucuna pruriens) technology by smallholder farmers in some areas in the derived savanna of West Africa. What is not known, however, is the extent to which the establishment and N contribution of mucuna in these areas depend on symbiotic properties such as effective nodulation and mycorrhizal infection. Short term surveys carried out in 34 farmer's arable fields located in four different sites in the derived savanna, southern Benin, West Africa, together with results of greenhouse and field experiments showed that mycorrhizal infection rate of mucuma ranged from 2 to 31% and correlated positively with nodulation and shoot dry matter production. Nodulation occurred in 79% of the fields with numbers of nodules ranging from 0 to 135 plant^–1. Mucuna responded both to inoculation and N fertilizer in degraded soils but growth response depended on the rhizobia strains and mucuna varieties. Mucuna accumulated in 12 weeks about 313 kg N ha^–1 as either a sole crop or 166 kg N ha^–1 when mixed/intercropped with maize, respectively. Across all cropping systems it derived an average of 70% of its N from atmospheric N₂ (estimates made by the ¹⁵N isotope dilution method), representing 167 kg N ha^–1 per 12 weeks in the field. Mucuna interplanted with maize obtained a greater proportion of its nitrogen (74%) from fixation than did mucuna grown alone (66%) suggesting that competition for soil N influences the proportion of nitrogen fixed by mucuna. The total amount of N₂ fixed per hectare was, however, reduced significantly by intercropping mucuna with maize. A preceding mucuna crop provided a maize yield equivalent to 120 kg N kg ha^–1 of inorganic N fertilizer.