Does deep soil N availability sustain long‐term ecosystem responses to elevated CO2?

A scrub‐oak woodland has maintained higher aboveground biomass accumulation after 11 years of atmospheric CO₂ enrichment (ambient +350 μmol CO₂ mol^?1), despite the expectation of strong nitrogen (N) limitation at the site. We hypothesized that changes in plant available N and exploitation of deep sources of inorganic N in soils have sustained greater growth at elevated CO₂. We employed a suite of assays performed in the sixth and 11th year of a CO₂ enrichment experiment designed to assess soil N dynamics and N availability in the entire soil profile. In the 11th year, we found no differences in gross N flux, but significantly greater microbial respiration (P≤0.01) at elevated CO₂. Elevated CO₂ lowered extractable inorganic N concentrations (P=0.096) considering the whole soil profile (0–190 cm). Conversely, potential net N mineralization, although not significant in considering the entire profile (P=0.460), tended to be greater at elevated CO₂. Ion‐exchange resins placed in the soil profile for approximately 1 year revealed that potential N availability at the water table was almost 3 × greater than found elsewhere in the profile, and we found direct evidence using a ¹⁵N tracer study that plants took up N from the water table. Increased microbial respiration and shorter mean residence times of inorganic N at shallower depths suggests that enhanced SOM decomposition may promote a sustained supply of inorganic N at elevated CO₂. Deep soil N availability at the water table is considerable, and provides a readily available source of N for plant uptake. Increased plant growth at elevated CO₂ in this ecosystem may be sustained through greater inorganic N supply from shallow soils and N uptake from deep soil.     

Effects of elevated atmospheric CO2 on soil microbiota in calcareous grassland

Microbial responses to three years of CO₂ enrichment (600 μL L^–1) in the field were investigated in calcareous grassland. Microbial biomass carbon (C) and soil organic C and nitrogen (N) were not significantly influenced by elevated CO₂. Microbial C:N ratios significantly decreased under elevated CO₂ (– 15%, P = 0.01) and microbial N increased by + 18% (P = 0.04). Soil basal respiration was significantly increased on one out of 7 sampling dates (+ 14%, P = 0.03; December of the third year of treatment), whereas the metabolic quotient for CO₂ (qCO₂ = basal respiration/microbial C) did not exhibit any significant differences between CO₂ treatments. Also no responses of microbial activity and biomass were found in a complementary greenhouse study where intact grassland turfs taken from the field site were factorially treated with elevated CO₂ and phosphorus (P) fertilizer (1 g P m^–2 y^–1). Previously reported C balance calculations showed that in the ecosystem investigated growing season soil C inputs were strongly enhanced under elevated CO₂. It is hypothesized that the absence of microbial responses to these enhanced soil C fluxes originated from mineral nutrient limitations of microbial processes. Laboratory incubations showed that short-term microbial growth (one week) was strongly limited by N availability, whereas P was not limiting in this soil. The absence of large effects of elevated CO₂ on microbial activity or biomass in such nutrient-poor natural ecosystems is in marked contrast to previously published large and short-term microbial responses to CO₂ enrichment which were found in fertilized or disturbed systems. It is speculated that the absence of such responses in undisturbed natural ecosystems in which mineral nutrient cycles have equilibrated over longer periods of time is caused by mineral nutrient limitations which are ineffective in disturbed or fertilized systems and that therefore microbial responses to elevated CO₂ must be studied in natural, undisturbed systems.     

Growth of Eastern Cottonwoods (Populus deltoides) in elevated [CO2] stimulates stand-level respiration and rhizodeposition of carbohydrates, accelerates soil nutrient depletion, yet stimulates above- and belowground biomass production

We took advantage of the distinctive system‐level measurement capabilities of the Biosphere 2 Laboratory (B2L) to examine the effects of prolonged exposure to elevated [CO₂] on carbon flux dynamics, above‐ and belowground biomass changes, and soil carbon and nutrient capital in plantation forest stands over 4 years. Annually coppiced stands of eastern cottonwoods (Populus deltoides) were grown under ambient (400 ppm) and two levels of elevated (800 and 1200 ppm) atmospheric [CO₂] in carbon and N‐replete soils of the Intensive Forestry Mesocosm in the B2L. The large semiclosed space of B2L uniquely enabled precise CO₂ exchange measurements at the near ecosystem scale. Highly controllable climatic conditions within B2L also allowed for reproducible examination of CO₂ exchange under different scales in space and time. Elevated [CO₂] significantly stimulated whole‐system maximum net CO₂ influx by an average of 21% and 83% in years 3 and 4 of the experiment. Over the 4‐year experiment, cumulative belowground, foliar, and total aboveground biomass increased in both elevated [CO₂] treatments. After 2 years of growth at elevated [CO₂], early season stand respiration was decoupled from CO₂ influx aboveground, presumably because of accelerated fine root production from stored carbohydrates in the coppiced system prior to canopy development and to the increased soil carbohydrate status under elevated [CO₂] treatments. Soil respiration was stimulated by elevated [CO₂] whether measured at the system level in the undisturbed soil block, by soil collars in situ, or by substrate‐induced respiration in vitro. Elevated [CO₂] accelerated depletion of soil nutrients, phosphorus, calcium and potassium, after 3 years of growth, litter removal, and coppicing, especially in the upper soil profile, although total N showed no change. Enhancement of above‐ and belowground biomass production by elevated [CO₂] accelerated carbon cycling through the coppiced system and did not sequester additional carbon in the soil.     

Potential impacts of climate change on nitrogen transformations and greenhouse gas fluxes in forests: a soil transfer study

Relatively little research has been conducted on how climate change may affect the structure and function of arid to semiarid ecosystems of the American Southwest. Along the slopes of the San Francisco Peaks of Arizona, USA, I transferred intact soil cores from a spruce‐fir to a ponderosa pine forest 730 m lower in elevation to assess the potential impacts of climate change on soil N cycling and trace gas fluxes. The low‐elevation site has a mean annual soil temperature about 2.5°C higher than the high‐elevation site. Net rates of N transformations and trace gas fluxes were measured in high‐elevation soil cores incubated in situ and soil cores transferred to the low‐elevation site. Over a 13‐month period, volumetric soil water content was similar in transferred soil cores relative to soil cores incubated in situ. Net N mineralization and nitrification increased over 80% in transferred soil cores compared with in situ soil cores. Soil transfer significantly increased net CO₂ efflux (120%) and net CH₄ consumption (90%) relative to fluxes of these gases from soil cores incubated in situ. Soil net N₂O fluxes were relatively low and were not generally altered by soil transfer. Although the soil microbial biomass as a whole decreased in transferred soil cores compared with in situ soil cores after the incubation period, active bacterial biomass increased. Transferring soil cores from the low‐elevation to the high‐elevation site (i.e. simulated global cooling) commonly, but not consistently, resulted in the opposite effects on soil pools and processes. In general, soil containment (root trenching) did not significantly affect soil measurements. My results suggest that small increases in mean annual temperature can have large impacts on soil N cycling, soil–atmosphere trace gas exchanges, and soil microbial communities even in ecosystems where water availability is a major limiting resource.     

Winter soil frost conditions in boreal forests control growing season soil CO2 concentration and its atmospheric exchange

The impact of changes in winter soil frost regime on soil CO₂ concentration and its atmospheric exchange in a boreal Norway spruce forest was investigated using a field‐scale soil frost manipulation experiment. The experiment comprised three treatments: deep soil frost, shallow soil frost and control plots (n= 3). Winter soil temperatures and soil frost distribution were significantly altered by the different treatments. The average soil CO₂ concentrations during the growing season were significantly lower in plots with deep soil frost than in plots with shallow soil frost. The average CO₂ soil–atmosphere exchange rate exhibited the same pattern, and differences in soil respiration rates among the treatments were statistically significant. Both the variation in soil CO₂ concentration and the CO₂ soil–atmosphere exchange rate could statistically be explained by the differences in the maximum soil frost depth during the previous winter. A response model for growing season soil respiration rates suggests that every 1 cm change in winter soil frost depth will change the emission rates by ca. 0.01 g CO₂ m^?2 day^?1, corresponding to 0.2–0.5% of the estimated net ecosystem productivity (NEP). This suggests that the soil frost regime has a significant influence on the C balance of the system, because interannual variations in soil frost up to 60 cm have been recorded at the site. We conclude that winter climate conditions can be important in controlling C balances in northern terrestrial ecosystems, and also that indirect effects of the winter season must be taken into account, because these can affect the prevailing conditions during the growing season.     

Nutrient Input and Carbon and Microbial Dynamics in an Ombrotrophic Bog

Slow rates of plant production and decomposition in ombrotrophic bogs are believed to be partially the result of low nutrient availability. To test the effect of nutrient availability on decomposition, carbon dioxide (CO₂) flux dynamics, microbial biomass, and nutrients, we added nitrogen (N) with phosphorus (P) and potassium (K), to prevent limitation of the latter 2 nutrients, over 2 growing seasons to plots at Mer Bleue peatland, Ontario, Canada. After the first growing season, increasing N fertilization (with constant P and K) decreased in vitro CO₂ production potential and increased microbial biomass measured with a chloroform fumigation-extraction technique in the upper peat profile, while by the end of the second season, CO₂ production potential was increased in response to N plus PK treatment, presumably due to more easily decomposable newly formed plant material. In situ CO₂ fluxes measured using chamber-techniques over the second year corroborated this presumption, with greater photosynthetic CO₂ uptake and ecosystem respiration (ER) during high N plus PK treatments. The more efficient microbial community, with slower CO₂ production potential and larger biomass, after the first year was characterized by larger fungal biomass measured with signature phospholipid fatty acids. The majority of N was likely quickly sequestered by the vegetation and transferred to dissolved organic forms and microbial biomass in the upper parts of the peat profile, while additional P relative to controls was distributed throughout the profile, implying that the vegetation at the site was N limited. However, in situ CO₂ flux data suggested the possibility of P or NPK limitation. We hypothesize that nutrient deposition may lead to enhanced C uptake by altering the microbial community and decomposition, however this pattern disappears through subsequent changes in the vegetation and production of more readily decomposable plant tissues.     

Glycine uptake in heath plants and soil microbes responds to elevated temperature, CO2 and drought

Temperate terrestrial ecosystems are currently exposed to climatic and air quality changes with increased atmospheric CO₂, increased temperature and prolonged droughts. The responses of natural ecosystems to these changes are focus for research, due to the potential feedbacks to the climate. We here present results from a field experiment in which the effects of these three climate change factors are investigated solely and in all combinations at a temperate heath dominated by heather (Calluna vulgaris) and wavy hair-grass (Deschampsia flexuosa).Climate induced increases in plant production may increase plant root exudation of dissolved organic compounds such as amino acids, and the release of amino acids during decomposition of organic matter. Such free amino acids in soil serve as substrates for soil microorganisms and are also acquired as nutrients directly by plants. We investigated the magnitude of the response to the potential climate change treatments on uptake of organic nitrogen in an in situ pulse labelling experiment with ¹⁵N¹³C₂-labelled glycine (amino acid) injected into the soil.In situ root nitrogen acquisition by grasses responded significantly to the climate change treatments, with larger ¹⁵N uptake in response to warming and elevated CO₂ but not additively when the treatments were combined. Also, a larger grass leaf biomass in the combined T and CO₂ treatment than in individual treatments suggest that responses to combined climate change factors cannot be predicted from the responses to single factors treatments.The soil microbes were superior to plants in the short-term competition for the added glycine, as indicated by an 18 times larger ¹⁵N recovery in the microbial biomass compared to the plant biomass. The soil microbes acquired glycine largely as an intact compound (87%), with no effects of the multi factorial climate change treatment through one year.     

Net mineralization of N at deeper soil depths as a potential mechanism for sustained forest production under elevated [CO2]

Elevated atmospheric carbon dioxide concentrations [CO₂] is projected to increase forest production, which could increase ecosystem carbon (C) storage. This study contributes to our broad goal of understanding the causes and consequences of increased fine‐root production and mortality under elevated [CO₂] by examining potential gross nitrogen (N) cycling rates throughout the soil profile. Our study was conducted in a CO₂‐enriched sweetgum (Liquidambar styraciflua L.) plantation in Oak Ridge, TN, USA. We used ¹⁵N isotope pool dilution methodology to measure potential gross N cycling rates in laboratory incubations of soil from four depth increments to 60 cm. Our objectives were twofold: (1) to determine whether N is available for root acquisition in deeper soil and (2) to determine whether elevated [CO₂], which has increased inputs of labile C resulting from greater fine‐root mortality at depth, has altered N cycling rates. Although gross N fluxes declined with soil depth, we found that N is potentially available for roots to access, especially below 15 cm depth where rates of microbial consumption of mineral N were reduced relative to production. Overall, up to 60% of potential gross N mineralization and 100% of potential net N mineralization occurred below 15 cm depth at this site. This finding was supported by in situ measurements from ion‐exchange resins, where total inorganic N availability at 55 cm depth was equal to or greater than N availability at 15 cm depth. While it is likely that trees grown under elevated [CO₂] are accessing a larger pool of inorganic N by mining deeper soil, we found no effect of elevated [CO₂] on potential gross or net N cycling rates. Thus, increased root exploration of the soil volume under elevated [CO₂] may be more important than changes in potential gross N cycling rates in sustaining forest responses to rising atmospheric CO₂.     

Living roots magnify the response of soil organic carbon decomposition to temperature in temperate grassland

Increasing atmospheric carbon dioxide (CO₂) concentration is both a strong driver of primary productivity and widely believed to be the principal cause of recent increases in global temperature. Soils are the largest store of the world's terrestrial C. Consequently, many investigations have attempted to mechanistically understand how microbial mineralisation of soil organic carbon (SOC) to CO₂ will be affected by projected increases in temperature. Most have attempted this in the absence of plants as the flux of CO₂ from root and rhizomicrobial respiration in intact plant‐soil systems confounds interpretation of measurements. We compared the effect of a small increase in temperature on respiration from soils without recent plant C with the effect on intact grass swards. We found that for 48 weeks, before acclimation occurred, an experimental 3 °C increase in sward temperature gave rise to a 50% increase in below ground respiration (ca. 0.4 kg C m^?2; Q₁₀ = 3.5), whereas mineralisation of older SOC without plants increased with a Q₁₀ of only 1.7 when subject to increases in ambient soil temperature. Subsequent ¹⁴C dating of respired CO₂ indicated that the presence of plants in swards more than doubled the effect of warming on the rate of mineralisation of SOC with an estimated mean C age of ca. 8 years or older relative to incubated soils without recent plant inputs. These results not only illustrate the formidable complexity of mechanisms controlling C fluxes in soils but also suggest that the dual biological and physical effects of CO₂ on primary productivity and global temperature have the potential to synergistically increase the mineralisation of existing soil C.     

Elevated atmospheric CO2 affects decomposition of Festuca vivipara (L.) Sm. litter and roots in experiments simulating environmental change in two contrasting arctic ecosystems

Mass loss, together with nitrogen and carbon loss, from above-ground material and roots of Festuca vivipara were followed for 13 months in a high Arctic polar semi-desert and a low Arctic tree-line dwarf shrub heath. Festuca vivipara for the study was obtained from plants cultivated at two different CO₂ concentrations (350 and 500 μL L^–1) in controlled environment chambers in the UK. Each of the four resource types (shoots or roots from plants grown in elevated or ambient CO₂ concentrations) was subsequently placed in an experiment simulating aspects of environmental change in each Arctic ecosystem. Air, litter and soil temperatures were increased using open-topped polythene tents at both sites, and a 58% increase in summer precipitation was simulated at the high Arctic site. Mass loss was greatest at the low Arctic site, and from the shoot material, rather than the roots. Shoots grown under an elevated CO₂ concentration decomposed more slowly at the high Arctic site, and more quickly at the low Arctic one, than shoots grown at ambient CO₂. After 13 months, greater amounts of C and N remained in above-ground litter from plants grown under elevated, rather than ambient, CO₂ at the polar semi-desert site, although lower amounts of C remained in elevated CO₂ litter at the low Arctic ecosystem. In the high Arctic, roots grown in the 500 μL L^–1 CO₂ concentration decomposed significantly more slowly than below-ground material derived from the ambient CO₂ chambers. Elevated CO₂ concentrations significantly increased the inital C:N ratio, % soluble carbohydrates and α-cellulose content, and significantly decreased the inital N content, of the above-ground material compared to that derived from the ambient treatment. Initially, the C:N ratio and percentage N were similar in both sets of roots derived from the two different CO₂ treatments, but soluble carbohydrate and α-cellulose concentrations were higher, and percentage lignin lower, in the elevated CO₂ treatments.The tent treatments significantly retarded shoot decomposition in both ecosystems, probably because of lower litter bag moisture contents, although the additional precipitation treatment had no effect on mass loss from the above-ground material. The results suggest that neither additional summer precipitation (up to 58%), nor soil temperature increase of 1 °C, which may occur by the end of the next century as an effect of a predicted 4 °C rise in air temperature, had an appreciable effect on root decomposition in the short term in a high Arctic soil. However, at the low Arctic site, greater root decomposition, and a lower pool of root N remaining, were observed where soil temperature was increased by 2 °C in response to a 4 °C rise in air temperature. These results suggest that decomposition below-ground in this ecosystem would increase as an effect of predicted climate change. These data also show that there is a difference in the initial results of decomposition processes between the two Arctic ecosystems in response to simulated environmental change.     

Development of nitrogen mineralisation gradients through surface soil depth and their influence on surface soil pH

Following mixing of the surface soil to about 7.5 cm depth in the field, soil layers (0–2.5, 2.5–5, 5–10 and 10–15 cm) were separately incubated in the laboratory to determine the rate of development of net N mineralisation gradients through surface soil depth under fallow, wheat and subterranean clover plots. Gradients in net N mineralisation were compared with those observed in the field, and their contribution to the observed pH changes was investigated.Heterotrophic activity, and thus net N mineralisation, decreased only slightly with depth immediately after soil mixing. This pattern persisted over time in soil layers sampled from fallow plots. In contrast, within 1 growing season after soil mixing, heterotrophic activity and net N mineralisation decreased significantly with depth in soil sampled from wheat and clover plots. In 0–15 cm soil sampled from under senescing plants, 32–38% of CO₂-C produced and net N mineralised originated from the surface 2.5 cm, while 52–56% originated from the surface 5 cm of soil. This resulted from an increase of pH and organic substrate concentration within the surface 2.5 cm of soil following plant residue return. Limitations of the in situ measurement of net N mineralisation in fallow soil was identified.Laboratory incubation studies showed that since most net N mineralisation occurred within the surface 2.5 cm of soil under senescing plants, nitrification and acidification were also concentrated at this depth. Despite this, compared to fallow soil, high potential acidification rates of 0–2.5 cm soil under senescing plants were not realised in the field due to the exposure to prolonged dry periods and moist-dry cycles. As a consequence, in the field the large magnitude of surface soil pH gradient which resulted from the return of alkaline plant residues was maintained over summer and autumn.     

Transferring soils from high- to low-elevation forests increases nitrogen cycling rates: climate change implications

We assessed the potential impact of global warming resulting from a doubling of preindustrial atmospheric CO₂ on soil net N transformations by transferring intact soil cores (0–15 cm) from a high-elevation old-growth forest to a forest about 800 m lower in elevation in the central Oregon Cascade Mountains, USA. The lower elevation site had mean annual air and soil (10-cm mineral soil depth) temperatures about 2.4 and 3.9 °C higher than the high-elevation site, respectively. Annual rates of soil net N mineralization and nitrification more than doubled in soil transferred to the low-elevation site (17.2–36.0 kg N ha^–1 and 5.0–10.7 kg NO₃^––N ha^–1, respectively). Leaching of inorganic N from the surface soil (in the absence of plant uptake) also increased. The reciprocal treatment (transferring soil cores from the low- to the high-elevation site) resulted in decreases of about 70, 80, and 65% in annual rates of net N mineralization, nitrification, and inorganic N leaching, respectively. Laboratory incubations of soils under conditions of similar temperature and soil water potential suggest that the quality of soil organic matter is higher at the high-elevation site. Similar in situ rates of soil net N transformations between the two sites occurred because the lower temperature counteracts the effects of greater substrate quantity and quality at the high elevation site. Our results support the hypothesis that high-elevation, old-growth forest soils in the central Cascades have higher C and N storage than their low-elevation analogues primarily because low temperatures limit net C and N mineralization rates at higher elevations.     

Effects of Fertilization on Soil CO₂ Efflux in Chinese Hickory (<i>Carya cathayensis</i>) Stand

Chinese hickory (Carya cathayensis Sarg.) is a popular nut tree in China, but there is little information about the influences of fertilization on soil CO₂ efflux and soil microbial biomass. This study evaluated the short-term effects of different fertilizer applications on soil CO₂ efflux and soil microbial biomass in Chinese hickory stands. Four fertilizer treatments were established: control (CK, no fertilizer), inorganic fertilizer (IF), organic fertilizer (OF), and equal parts organic and inorganic N fertilizers (OIF). A field experiment was conducted to measure soil CO₂ effluxes using closed chamber and gas chromatography techniques. Regardless of the fertilization practices, soil CO₂ effluxes of all the treatments showed a similar temporal pattern, with the highest value in summer and the lowest in winter. The mean annual soil CO₂ efflux in the IF treatment was significantly higher than that in the CK, OIF, and OF treatments. There was no significant difference in soil CO₂ efflux between the OIF, OF, and CK treatments. Soil CO₂ effluxes were significantly affected by soil temperature. Soil dissolved organic carbon (DOC) was positively correlated with soil CO₂ efflux only in the CK treatment. Regression analysis, including soil temperature, moisture, and DOC, showed that soil temperature was the primary factor influencing soil CO₂ effluxes. Both OF and OIF treatments increased concentrations of soil microbial biomass carbon (MBC) and microbial biomass nitrogen (MBN), but decreased the ratio of MBC:MBN. These results reveal that applying organic fertilizer, either alone or combined with inorganic fertilizer, may be the optimal strategy for mitigating soil CO₂ emission and improving soil quality in Chinese hickory stands.     

Responses of CO2, N2O and CH4 fluxes between atmosphere and forest soil to changes in multiple environmental conditions

To investigate the effects of multiple environmental conditions on greenhouse gas (CO₂, N₂O, CH₄) fluxes, we transferred three soil monoliths from Masson pine forest (PF) or coniferous and broadleaved mixed forest (MF) at Jigongshan to corresponding forest type at Dinghushan. Greenhouse gas fluxes at the in situ (Jigongshan), transported and ambient (Dinghushan) soil monoliths were measured using static chambers. When the transported soil monoliths experienced the external environmental factors (temperature, precipitation and nitrogen deposition) at Dinghushan, its annual soil CO₂ emissions were 54% in PF and 60% in MF higher than those from the respective in situ treatment. Annual soil N₂O emissions were 45% in PF and 44% in MF higher than those from the respective in situ treatment. There were no significant differences in annual soil CO₂ or N₂O emissions between the transported and ambient treatments. However, annual CH₄ uptake by the transported soil monoliths in PF or MF was not significantly different from that at the respective in situ treatment, and was significantly lower than that at the respective ambient treatment. Therefore, external environmental factors were the major drivers of soil CO₂ and N₂O emissions, while soil was the dominant controller of soil CH₄ uptake. We further tested the results by developing simple empirical models using the observed fluxes of CO₂ and N₂O from the in situ treatment and found that the empirical models can explain about 90% for CO₂ and 40% for N₂O of the observed variations at the transported treatment. Results from this study suggest that the different responses of soil CO₂, N₂O, CH₄ fluxes to changes in multiple environmental conditions need to be considered in global change study.     

Soil pCO2, soil respiration,and root activity in CO2-fumigated and nitrogen-fertilized ponderosa pine

The purpose of this paper is to describe the effects of CO₂ and N treatments on soil pCO₂, calculated CO₂ efflux, root biomass and soil carbon in open-top chambers planted with Pinus ponderosa seedlings. Based upon the literature, it was hypothesized that both elevated CO₂ and N would cause increased root biomass which would in turn cause increases in both total soil CO₂ efflux and microbial respiration. This hypothesis was only supported in part: both CO₂ and N treatments caused significant increases in root biomass, soil pCO₂, and calculated CO₂ efflux, but there were no differences in soil microbial respiration measured in the laboratory. Both correlative and quantitative comparisons of CO₂ efflux rates indicated that microbial respiration contributes little to total soil CO₂ efflux in the field. Measurements of soil pCO₂ and calculated CO₂ efflux provided inexpensive, non-invasive, and relatively sensitive indices of belowground response to CO₂ and N treatments.     

Soil–atmosphere exchange of greenhouse gases in a Eucalyptus marginata woodland, a clover-grass pasture, and Pinus radiata and Eucalyptus globulus plantations

Soils provide the largest terrestrial carbon store, the largest atmospheric CO₂ source, the largest terrestrial N₂O source and the largest terrestrial CH₄ sink, as mediated through root and soil microbial processes. A change in land use or management can alter these soil processes such that net greenhouse gas exchange may increase or decrease. We measured soil–atmosphere exchange of CO₂, N₂O and CH₄ in four adjacent land‐use systems (native eucalypt woodland, clover‐grass pasture, Pinus radiata and Eucalyptus globulus plantation) for short, but continuous, periods between October 2005 and June 2006 using an automated trace gas measurement system near Albany in southwest Western Australia. Mean N₂O emission in the pasture was 26.6 μg N m⁻² h⁻¹, significantly greater than in the natural and managed forests (< 2.0 μg N m⁻² h⁻¹). N₂O emission from pasture soil increased after rainfall events (up to 100 μg N m⁻² h⁻¹) and as soil water content increased into winter, whereas no soil water response was detected in the forest systems. Gross nitrification through ¹⁵N isotope dilution in all land‐use systems was small at water holding capacity < 30%, and under optimum soil water conditions gross nitrification ranged between < 0.1 and 1.0 mg N kg⁻¹ h⁻¹, being least in the native woodland/eucalypt plantation < pine plantation < pasture. Forest soils were a constant CH₄ sink, up to −20 μg C m⁻² h⁻¹ in the native woodland. Pasture soil was an occasional CH₄ source, but weak CH₄ sink overall (−3 μg C m⁻² h⁻¹). There were no strong correlations (R < 0.4) between CH₄ flux and soil moisture or temperature. Soil CO₂ emissions (35–55 mg C m⁻² h⁻¹) correlated with soil water content (R < 0.5) in all but the E. globulus plantation. Soil N₂O emissions from improved pastures can be considerable and comparable with intensively managed, irrigated and fertilised dairy pastures. In all land uses, soil N₂O emissions exceeded soil CH₄ uptake on a carbon dioxide equivalent basis. Overall, afforestation of improved pastures (i) decreases soil N₂O emissions and (ii) increases soil CH₄ uptake.     

Nutrient relations in calcareous grassland under elevated CO2

Plant nutrient responses to 4 years of CO₂ enrichment were investigated in situ in calcareous grassland. Beginning in year 2, plant aboveground C:N ratios were increased by 9% to 22% at elevated CO₂ (P < 0.01), depending on year. Total amounts of N removed in biomass harvests during the first 4 years were not affected by elevated CO₂ (19.9 ± 1.3 and 21.1 ± 1.3 g N m⁻² at ambient and elevated CO₂), indicating that the observed plant biomass increases were solely attained by dilution of nutrients. Total aboveground P and tissue N:P ratios also were not altered by CO₂ enrichment (12.5 ± 2 g N g⁻¹ P in both treatments). In contrast to non-legumes (>98% of community aboveground biomass), legume C/N was not reduced at elevated CO₂ and legume N:P was slightly increased. We attribute the less reduced N concentration in legumes at elevated CO₂ to the fact that virtually all legume N originated from symbiotic N₂ fixation (%N_dfa ≈ 90%), and thus legume growth was not limited by soil N. While total plant N was not affected by elevated CO₂, microbial N pools increased by +18% under CO₂ enrichment (P = 0.04) and plant available soil N decreased. Hence, there was a net increase in the overall biotic N pool, largely due increases in the microbial N pool. In order to assess the effects of legumes for ecosystem CO₂ responses and to estimate the degree to which plant growth was P-limited, two greenhouse experiments were conducted, using firstly undisturbed grassland monoliths from the field site, and secondly designed `microcosm' communities on natural soil. Half the microcosms were planted with legumes and half were planted without. Both monoliths and microcosms were exposed to elevated CO₂ and P fertilization in a factored design. After two seasons, plant N pools in both unfertilized monoliths and microcosm communities were unaffected by CO₂ enrichment, similar to what was found in the field. However, when P was added total plant N pools increased at elevated CO₂. This community-level effect originated almost solely from legume stimulation. The results suggest a complex interaction between atmospheric CO₂ concentrations, N and P supply. Overall ecosystem productivity is N-limited, whereas CO₂ effects on legume growth and their N₂ fixation are limited by P. Received: 12 July 1997 / Accepted: 15 April 1998     

Effects of elevated CO2 on nutrient cycling in a sweetgum plantation

The effects of elevated CO₂ on nutrient cycling and selected belowground processes in the closed-canopy sweetgum plantation were assessed as part of a free-air CO₂ enrichment (FACE) experiment at Oak Ridge, Tennessee. We hypothesized that nitrogen (N) constraints to growth response to elevated CO₂ would be mitigated primarily by reduced tissue concentrations (resulting in increased biomass production per unit uptake) rather than increased uptake. Conversely, we hypothesized that the constraints of other nutrients to growth response to elevated CO₂ would be mitigated primarily by increased uptake because of adequate soil supplies. The first hypothesis was not supported: although elevated CO₂ caused reduced foliar N concentrations, it also resulted in increased uptake and requirement of N, primarily because of greater root turnover. The additional N uptake with elevated CO₂ constituted between 10 and 40% of the estimated soil mineralizeable N pool. The second hypothesis was largely supported: elevated CO₂ had no significant effects on tissue concentrations of P, K, Ca, or Mg and caused significantly increased uptake and requirement of K, Ca, and Mg. Soil exchangeable pools of these nutrients are large and should pose no constraint to continued growth responses. Elevated CO₂ also caused increased microbial biomass, reduced N leaching and increased P leaching from O horizons (measured by resin lysimeters), reduced soil solution NH ₄ ⁺ , SO ₄ ^2– , and Ca²⁺ concentrations, and increased soil solution pH. There were no statistically significant treatment effects on soil nutrient availability as measured by resin capsules, resin stakes, or in situ incubations. Despite significantly lower litterfall N concentrations in the elevated CO₂ treatment, there were no significant treatment effects on translocation or forest floor biomass or nutrient contents. There were also no significant treatment effects on the rate of decomposition of fine roots. In general, the effects of elevated CO₂ on nutrient cycling in this study were not large; future constraints on growth responses imposed by N limitations will depend on changes in N demand, atmospheric N deposition, and soil mineralization rates.     

Biochemical properties of soils of undisturbed and disturbed mangrove forests of South Andaman (India)

Studies on soil quality of mangrove forests would be of immense use in minimizing soil degradation and in adopting strategies for soil management at degraded sites. Among the various parameters of soil quality, biological and biochemical soil properties are very sensitive to environmental stress and provide rapid and accurate estimates on changes in quality of soils subjected to degradation. In this study, we determined the general and specific biochemical characteristics of soils (0-30 cm) of inter-tidal areas of 10 undisturbed mangrove forest sites of S. Andaman, India. In order to determine the effects of disturbance, soils from the inter-tidal areas of 10 disturbed mangrove forest sites were also included in the study. The general biochemical properties included all the variables directly related to microbial activity and the specific biochemical parameters included the activities of extracellular hydrolytic enzymes that are involved in the carbon, nitrogen, sulfur and phosphorus cycles in soil. The pH, clay, cation exchange capacity, Al₂O₃ and Fe₂O₃ levels exhibited minimum variation between the disturbed and undisturbed sites. In contrast, organic C, total N, Bray P and K levels exhibited marked variation between the sites and were considerably lower at the disturbed sites. The study also revealed marked reductions in microbial biomass and activity at the disturbed sites. In comparison to the undisturbed sites, the levels of all the general biochemical parameters viz., microbial biomass C, microbial biomass N, N flush, basal respiration, metabolic quotient (qCO₂), ATP, N mineralization rates and the activities of dehydrogenase and catalase were considerably lower at the disturbed sites. Similarly, drastic reductions in the activities of phosphomonoesterase, phosphodiesterase, ß-g1ucosidase, urease, BAA-protease, casein-protease, arylsulfatase, invertase and carboxymethylcellulase occurred at the disturbed sites due mainly to significant reductions in organic matter/substrate levels. The data on CO₂ evolution, qCO₂ and ATP indicated the dominance of active individuals in the microbial communities of undisturbed soils and the ratios of biomass C:N, ATP:biomass C and ergosterol:biomass C ratios indicated low N availability and the possibility of fungi dominating over bacteria at both the mangrove sites. Significant and positive correlations between soil variables and biochemical properties suggested that the number and activity of soil microorganisms depend mainly on the quantity of mineralizable substrate and the availability of nutrients in these mangrove soils.     

Ponderosa Pine Responses to Elevated CO2and Nitrogen Fertilization

The effects of elevated CO₂ (ambient, +175, and +350 μl l⁻¹) and nitrogen fertilization (0, 100, and 200 kg N ha⁻¹ yr⁻¹ as ammonium sulfate) on C and N accumulations in biomass and soils planted with ponderosa pine (Pinus ponderosa Laws) over a 6-year study period are reported. Both nitrogen fertilization and elevated CO₂ caused increases in C and N contents of vegetation over the study period. The pattern of responses varied over time. Responses to CO₂ decreased in the +175 μl l⁻¹ and increased in the +350 μl l⁻¹ after the first year, whereas responses to N decreased after the first year and became non-significant by year six. Foliar N concentrations were lower and tree C:N ratios were higher with elevated CO₂ in the early years, but this was offset by the increases in biomass, resulting in substantial increases in N uptake with elevated CO₂. Nitrogen budget estimates showed that the major source of the N for unfertilized trees, with or without elevated CO₂, was likely the soil organic N pool. There were no effects of elevated CO₂ on soil C, but a significant decrease in soil N and an increase in soil C:N ratio in year six. Nitrogen fertilization had no significant effect on tree C:N ratios, foliar N concentrations, soil C content, soil N content, or soil C:N ratios. There were no significant interactions between CO₂ and N treatments, indicating that N fertilization had no effect on responses to CO₂ and that CO₂ treatments had no effect on responses to N fertilization. These results illustrate the importance of long-term studies involving more than one level of treatment to assess the effects of elevated CO₂.