Potassium limits potential growth of bog vegetation under elevated atmospheric CO2 and N deposition

The free air carbon dioxide enrichment (FACE) and N deposition experiments on four ombrotrophic bogs in Finland, Sweden, the Netherlands and Switzerland, revealed that after three years of treatment: (1) elevated atmospheric CO₂ concentration had no significant effect on the biomass growth of Sphagnum and vascular species; and (2) increased N deposition reduced Sphagnum growth, because it increased the cover of vascular plants and the tall moss Polytrichum strictum, while vascular plant biomass growth was not affected. This paper focuses on water chemistry, plant nutrient content, and litter decomposition rates. Potassium limitation, or low supply of K and P, may have prevented a significant increase of Sphagnum growth under elevated CO₂ and N deposition. Vascular plant growth under elevated CO₂ and N deposition was also limited by K, or by K in combination with P or N (N in CO₂ experiment). Elevated CO₂ and N deposition had no effect on decomposition rates of Sphagnum and vascular plant litter. Aside from a possible effect of N deposition on light competition between species, we expect that elevated atmospheric CO₂ and N deposition concentrations will not affect Sphagnum and vascular plant growth in bogs of north‐west Europe due to K‐, or K in combination with N‐ or P‐, limited growth. For the same reason we expect no effect of elevated CO₂ and N deposition on litter decomposition. Net primary production of raised ombrotrophic bogs that are at or close to steady state, is regulated by input of nutrients through atmospheric deposition. Therefore, we hypothesize that the expected increase of plant growth under elevated CO₂ and N deposition is diminished by current levels of K (and to some extent P and N) in atmospheric deposition.     

Effects of elevated CO2 and vascular plants on evapotranspiration in bog vegetation

We determined evapotranspiration in three experiments designed to study the effects of elevated CO₂ and increased N deposition on ombrotrophic bog vegetation. Two experiments used peat monoliths with intact bog vegetation in containers, with one experiment outdoors and the other in a greenhouse. A third experiment involved monocultures and mixtures of Sphagnum magellanicum and Eriophorum angustifolium in containers in the same greenhouse. To determine water use of the bog vegetation in July–August for each experiment and each year we measured water inputs and outputs from the containers. We studied the effects of elevated CO₂ and N supply on evapotranspiration in relation to vascular plant biomass and exposure of the moss surface (measured as height of the moss surface relative to the container edge). Elevated CO₂ reduced water use of the bog vegetation in all three experiments, but the CO₂ effect on evapotranspiration interacted with vascular plant biomass and exposure of the moss surface. Evapotranspiration in the outdoor experiment was largely determined by evaporation from the Sphagnum moss surface (as affected by exposure to wind) and less so by vascular plant transpiration. Nevertheless, elevated CO₂ significantly reduced evapotranspiration by 9–10% in the outdoor experiment. Vascular plants reduced evapotranspiration in the outdoor experiment, but increased water use in the greenhouse experiments. The relation between vascular plant abundance and evapotranspiration appears to depend on wind conditions; suggesting that vascular plants reduce water losses mainly by reducing wind speed at the moss surface. Sphagnum growth is very sensitive to changes in water level; low water availability can have deleterious effects. As a consequence, reduced evapotranspiration in summer, whether caused by elevated CO₂ or by small increases in vascular plant cover, is expected to favour Sphagnum growth in ombrotrophic bog vegetation.     

Turnover of labile and recalcitrant soil carbon differ in response to nitrate and ammonium deposition in an ombrotrophic peatland

The effects of 4 years of simulated nitrogen deposition, as nitrate (NO₃^?) and ammonium (NH₄⁺), on microbial carbon turnover were studied in an ombrotrophic peatland. We investigated the mineralization of simple forms of carbon using MicroResp^? measurements (a multiple substrate induced respiration technique) and the activities of four soil enzymes involved in the decomposition of more complex forms of carbon or in nutrient acquisition: N‐acetyl‐glucosaminidase (NAG), cellobiohydrolase (CBH), acid phosphatase (AP), and phenol oxidase (PO). The potential mineralization of labile forms of carbon was significantly enhanced at the higher N additions, especially with NH₄⁺ amendments, while potential enzyme activities involved in breakdown of more complex forms of carbon or nutrient acquisition decreased slightly (NAG and CBH) or remained unchanged (AP and PO) with N amendments. This study also showed the importance of distinguishing between NO₃^? and NH₄⁺ amendments, as their impact often differed. It is possible that the limited response on potential extracellular enzyme activity is due to other factors, such as limited exposure to the added N in the deeper soil or continued suboptimal functioning of the enzymes due to the low pH, possibly via the inhibitory effect of low phenol oxidase activity.     

Carbon balance of a European mountain bog at contrasting stages of regeneration

Carbon dioxide and methane (CH4) fluxes were measured in a cutover bog of the Jura Mountains (France) together with biotic and abiotic variables for two entire vegetation periods in order to compare the carbon balance of the bog at three stages of regeneration. Among all factors, air temperature and vegetation index (including leaf area of vascular plants, bryophyte density and bryophyte desiccation) were the two main determinants of ecosystem respiration and gross photosynthesis at light saturation. During 2004 and 2005, the vegetated plots acted as carbon sinks. Net carbon exchange ranged between 67 and 166 g C m(-2) yr(-1) for the Eriophorum-dominated plots and between 93 and 183 g C m(-2) yr(-1) for the Sphagnum-dominated plots. The bare peat plots represented a net carbon source (between -19 and -32 g C m(-2) yr(-1)). Methane fluxes accounted for a very small part of the total carbon efflux (< 2%). The recovery of vegetation in our naturally regenerating bog was beneficial for the carbon sequestration after the relatively short period of 20 yr.     

Contemporary carbon balance and late Holocene carbon accumulation in a northern peatland

Northern peatlands contain up to 25% of the world's soil carbon (C) and have an estimated annual exchange of CO₂‐C with the atmosphere of 0.1–0.5 Pg yr⁻¹ and of CH₄‐C of 10–25 Tg yr⁻¹. Despite this overall importance to the global C cycle, there have been few, if any, complete multiyear annual C balances for these ecosystems. We report a 6‐year balance computed from continuous net ecosystem CO₂ exchange (NEE), regular instantaneous measurements of methane (CH₄) emissions, and export of dissolved organic C (DOC) from a northern ombrotrophic bog. From these observations, we have constructed complete seasonal and annual C balances, examined their seasonal and interannual variability, and compared the mean 6‐year contemporary C exchange with the apparent C accumulation for the last 3000 years obtained from C density and age‐depth profiles from two peat cores. The 6‐year mean NEE‐C and CH₄‐C exchange, and net DOC loss are −40.2±40.5 (±1 SD), 3.7±0.5, and 14.9±3.1 g m⁻² yr⁻¹, giving a 6‐year mean balance of −21.5±39.0 g m⁻² yr⁻¹ (where positive exchange is a loss of C from the ecosystem). NEE had the largest magnitude and variability of the components of the C balance, but DOC and CH₄ had similar proportional variabilities and their inclusion is essential to resolve the C balance. There are large interseasonal and interannual ranges to the exchanges due to variations in climatic conditions. We estimate from the largest and smallest seasonal exchanges, quasi‐maximum limits of the annual C balance between 50 and −105 g m⁻² yr⁻¹. The net C accumulation rate obtained from the two peatland cores for the interval 400–3000 bp (samples from the anoxic layer only) were 21.9±2.8 and 14.0±37.6 g m⁻² yr⁻¹, which are not significantly different from the 6‐year mean contemporary exchange.     

Elevated UV-B radiation alters fluxes of methane and carbon dioxide in peatland microcosms

Increases in solar ultraviolet‐B radiation (UV‐B; 280–320 nm) reaching the earth have been estimated to continue until 2050s in the boreal and subarctic regions with an abundant peatland cover. Peatlands are significant sinks for carbon dioxide (CO₂) and sources for methane (CH₄). To assess whether the future increases in UV‐B could affect the fluxes of CO₂ and CH₄ in peatlands via an impact on vegetation, we exposed peatland microcosms to modulated 30% supplementation of erythemally weighted UV‐B at an outdoor facility for one growing season. The experimental design included appropriate controls for UV‐A and ambient radiation. The UV‐B caused a significant reduction in gross photosynthesis, net ecosystem CO₂ exchange, and CH₄ emission of the peatland microcosms. These changes in the carbon gas cycling can be partly explained by UV‐B‐induced morphological changes in Eriophorum vaginatum which acts as a conduit for CH₄. Leaf cross section and the percentage of CH₄‐conducting aerenchymatous tissue in E. vaginatum were significantly reduced by UV‐B. Methanol‐extractable UV‐B absorbing compounds decreased under both UV‐B and UV‐A in Sphagnum angustifolium, and tended to accumulate under UV‐B in S. papillosum. Membrane permeability to magnesium (Mg) and calcium (Ca) ions was higher in UV‐B exposed S. angustifolium. Amount of chlorophyll and carotenoid pigments was increased by UV‐A in S. magellanicum. The observed changes in Sphagnum mosses did not coincide with those in carbon gas fluxes but occurred at the time of the highest UV intensity in the mid summer. Our findings indicate that increasing UV‐B may have more substantial effects on gas exchange in peatlands than previously thought.     

Postfire carbon balance in boreal bogs of Alberta, Canada

Boreal peatland ecosystems occupy about 3.5 million km² of the earth's land surface and store between 250 and 455 Pg of carbon (C) as peat. While northern hemisphere boreal peatlands have functioned as net sinks for atmospheric C since the most recent deglaciation, natural and anthropogenic disturbances, and most importantly wildfire, may compromise peatland C sinks. To examine the effects of fire on local and regional C sink strength, we focused on a 12 000 km² region near Wabasca, AB, Canada, where ombrotrophic Sphagnum‐dominated bogs cover 2280 km² that burn with a fire return interval of 123±26 years. We characterized annual C accumulation along a chronosequence of 10 bog sites, spanning 1–102 years‐since‐fire (in 2002). Immediately after fire, bogs represent a net C source of 8.9±8.4 mol m⁻² yr⁻¹. At about 13 years after fire, bogs switch from net C sources to net C sinks, mainly because of recovery of the moss and shrub layers. Subsequently, black spruce biomass accumulation contributes to the net C sink, with fine root biomass accumulation peaking at 34 years after fire and aboveground biomass and coarse root accumulation peaking at 74 years after fire. The overall C sink strength peaks at 18.4 mol C m⁻² yr⁻¹ at 75 years after fire. As the tree biomass accumulation rate declines, the net C sink decreases to about 10 mol C m⁻² yr⁻¹ at 100 years‐since‐fire. We estimate that across the Wabasca study region, bogs currently represent a C sink of 14.7±5.1 Gmol yr⁻¹. A decrease in the fire return interval to 61 years with no change in air temperature would convert the region's bogs to a net C source. An increase in nonwinter air temperature of 2 °C would decrease the regional C sink to 6.8±2.3 Gmol yr⁻¹. Under scenarios of predicted climate change, the current C sink status of Alberta bogs is likely to diminish to the point where these peatlands become net sources of atmospheric CO₂‐C.     

Patterns of nitrogen and sulfur accumulation and retention in ombrotrophic bogs, eastern Canada

Nitrogen (N) and sulfur (S) play important roles in peatlands, through their influence on plant production and peat decomposition rates and on redox reactions, respectively, and peatlands contain substantial stores of these two elements. Using peat N and S concentrations and dry bulk density and ²¹⁰Pb dating, we determined the rates of N and S accumulation over the past 150 years in hummock and hollow profiles from 23 ombrotrophic bogs in eastern Canada. Concentrations of N and S averaged 0.80% and 0.18%, respectively, generally increased with depth in the profile and there was a weak but significant correlation between N and S concentrations. Rates of N and S accumulation over the past 50–150 years ranged from 0.5 to 4.8 g N m^?2 yr^?1 and from 0.1 to 0.9 g S m^?2 yr^?1. There were significant but weak correlations between C, N and S accumulation rates over 50‐, 100‐ and 150‐year periods. Over the last 50 years, rates of S accumulation showed little differentiation between hummocks and hollows, whereas the pattern for N accumulation was more variable (hummock minus hollow rate ranged from ?1 to +1.5 g N m^?2 yr^?1), with hummocks generally having a larger N accumulation rate, correlated with the rate of carbon (C) accumulation. There was a modest but significant positive correlation between 50‐year rates of N accumulation and wet atmospheric deposition of N measured between 1990 and 1996, with accumulation rates about four times that of wet deposition. The difference between deposition and accumulation of N is attributed to organic N deposition, dry deposition and N₂ fixation. A weaker, but still significant, correlation was observed between 50‐year S accumulation and 1990–1996 wet atmospheric S deposition, with about 75% of the deposited S accumulating in the peat. A laboratory experiment with peat cores exposed to varying water table position and simulated N and S deposition, showed that on average 87% and 98% of the deposited NH₄⁺ and NO₃^?, respectively, and 58% of the deposited S were retained in the vegetation and unsaturated zone of the cores, supporting the results from the field study.     

Interannual variability in carbon dioxide fluxes and flux–climate relationships on grazed and ungrazed northern mixed-grass prairie

The annual carbon (C) budget of grasslands is highly dynamic, dependent on grazing history and on effects of interannual variability (IAV) in climate on carbon dioxide (CO₂) fluxes. Variability in climatic drivers may directly affect fluxes, but also may indirectly affect fluxes by altering the response of the biota to the environment, an effect termed ‘functional change’. We measured net ecosystem exchange of CO₂ (NEE) and its diurnal components, daytime ecosystem CO₂ exchange (P_D) and night‐time respiration (R_E), on grazed and ungrazed mixed‐grass prairie in North Dakota, USA, for five growing seasons. Our primary objective was to determine how climatic anomalies influence variability in CO₂ exchange. We used regression analysis to distinguish direct effects of IAV in climate on fluxes from functional change. Functional change was quantified as the improvement in regression on fitting a model in which slopes of flux–climate relationships vary among years rather than remain invariant. Functional change and direct effects of climatic variation together explained about 20% of variance in weekly means of NEE, P_D, and R_E. Functional change accounted for more than twice the variance in fluxes of direct effects of climatic variability. Grazing did not consistently influence the contribution of functional change to flux variability, but altered which environmental variable best explained year‐to‐year differences in flux–climate slopes, reduced IAV in seasonal means of fluxes, lessened the strength of flux–climate correlations, and increased NEE by reducing R_E relatively more than P_D. Most of these trends are consistent with the interpretation that grazing reduced the influence of plants on ecosystem fluxes. Because relationships between weekly values of fluxes and climatic regulators changed annually, year‐to‐year differences in the C balance of these ecosystems cannot be predicted from knowledge of IAV in climate alone.     

Towards an understanding of the relations between tree nutrition,nutrient cycling and environment

The paper contains a discussion of the interrelations between the sciences used by managers of forest land to improve their management, in particular with respect to the plant nutrient economy of the forest ecosystems. Both site studies and studies of nutrient cycling have been carried out for long periods without proper consideration of tree nutrition. Therefore these studies contributed less to the understanding of the role of nutrients as regulators of processes in ecosystems than might have been expected. This situation has improved, especially within the last decade. In addition the necessity to manage forest land for environmental values as well as for forest yield requires new interdisciplinary approaches in the study of the roles of plant nutrients in the forest. Even more branches of biological and environmental sciences than those just mentioned must be involved.     

Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model

The Lund–Potsdam–Jena Dynamic Global Vegetation Model (LPJ) combines process‐based, large‐scale representations of terrestrial vegetation dynamics and land‐atmosphere carbon and water exchanges in a modular framework. Features include feedback through canopy conductance between photosynthesis and transpiration and interactive coupling between these ‘fast’ processes and other ecosystem processes including resource competition, tissue turnover, population dynamics, soil organic matter and litter dynamics and fire disturbance. Ten plants functional types (PFTs) are differentiated by physiological, morphological, phenological, bioclimatic and fire‐response attributes. Resource competition and differential responses to fire between PFTs influence their relative fractional cover from year to year. Photosynthesis, evapotranspiration and soil water dynamics are modelled on a daily time step, while vegetation structure and PFT population densities are updated annually. Simulations have been made over the industrial period both for specific sites where field measurements were available for model evaluation, and globally on a 0.5°° × 0.5°° grid. Modelled vegetation patterns are consistent with observations, including remotely sensed vegetation structure and phenology. Seasonal cycles of net ecosystem exchange and soil moisture compare well with local measurements. Global carbon exchange fields used as input to an atmospheric tracer transport model (TM2) provided a good fit to observed seasonal cycles of CO₂ concentration at all latitudes. Simulated inter‐annual variability of the global terrestrial carbon balance is in phase with and comparable in amplitude to observed variability in the growth rate of atmospheric CO₂. Global terrestrial carbon and water cycle parameters (pool sizes and fluxes) lie within their accepted ranges. The model is being used to study past, present and future terrestrial ecosystem dynamics, biochemical and biophysical interactions between ecosystems and the atmosphere, and as a component of coupled Earth system models.     

Physiological and ecological controls on carbon sequestering in terrestrial ecosystems

Carbon cycling processes in ecosystems are generally believed to be well understood. Carbon, hydrogen, oxygen and other essential elements are chemically converted from inorganic to organic compounds primarily in the process of photosynthesis. Secondary metabolic processes cycle carbon in and among organisms and carbon is ultimately released back to the environment as CO₂ by respiratory processes. Unfortunately, our understanding of this cycle was determined under the assumption that the primary inorganic form of C (CO₂ in the atmosphere) was relatively constant. With the emerging concensus that atmospheric carbon concentration is increasing, we must now reassess our understanding of the carbon cycle. How will plants, animals and decomposers respond to a doubling of carbon supply? Will biological productivity be accelerated? If plant productivity increases will a predictable percentage of the increase be accumulated as increased standing crop? Or, is it possible that doubling the availability of CO₂ will increase metabolic activity at all trophic levels resulting in no net increase in system standing crop? The purpose of this paper is to review evidence for physiological and growth responses of plants to carbon dioxide enhancement. Essentially no research has been completed on the ecological aspects of these questions. From this review, I conclude that accurate predictions of future ecosystem responses to increasing atmospheric carbon dioxide concentration are not possible without additional understanding of physiological and ecological mechanisms.     

Organic carbon cycling in Taylor Valley, Antarctica: quantifying soil reservoirs and soil respiration

Organic carbon reservoirs and respiration rates in soils have been calculated for most major biomes on Earth revealing patterns related to temperature, precipitation, and location. Yet data from one of the Earth's coldest, driest, and most southerly soil ecosystems, that of the McMurdo Dry Valleys of Antarctica, are currently not a part of this global database. In this paper, we present the first regional calculations of the soil organic carbon reservoirs in a dry valley ecosystem (Taylor Valley) and report measurements of CO₂ efflux from Antarctic soils. Our analyses indicate that, despite the absence of visible accumulations of organic matter in most of Taylor Valley's arid soils, this soil environment contained a significant percentage (up to 72%) of the seasonally unfrozen organic carbon reservoir in the terrestrial ecosystem. Field measurements of soil CO₂‐efflux in Taylor Valley soils were used to evaluate biotic respiration and averaged 0.10 ± 0.08 μmol CO₂ m^?2 s^?1. Laboratory soil microcosms suggested that this respiration rate was sensitive to increases in temperature, moisture, and carbon addition. Finally, a steady‐state calculation of the mean residence time for organic carbon in Taylor Valley soils was 23 years. Because this value contradicts all that is currently known about carbon cycling rates in the dry valleys, we suggest that the dry valley soil carbon dynamics is not steady state. Instead, we suggest that the dynamic is complex, with at least two (short‐ and long‐term) organic carbon reservoirs. We also suggest that organic carbon in the dry valley soil environment may be more important, and play a more active role in long‐term ecosystem processes, than previously believed.     

Exposure to an enriched CO2 atmosphere alters carbon assimilation and allocation in a pine forest ecosystem

We linked a leaf-level CO₂ assimilation model with a model that accounts for light attenuation in the canopy and measurements of sap-flux-based canopy conductance into a new canopy conductance-constrained carbon assimilation (4C-A) model. We estimated canopy CO₂ uptake (A_nC) at the Duke Forest free-air CO₂ enrichment (FACE) study. Rates of A_nC estimated from the 4C-A model agreed well with leaf gas exchange measurements (A_net) in both CO₂ treatments. Under ambient conditions, monthly sums of net CO₂ uptake by the canopy (A_nC) were 13% higher than estimates based on eddy-covariance and chamber measurements. Annual estimates of A_nC were only 3% higher than carbon (C) accumulations and losses estimated from ground-based measurements for the entire stand. The C budget for the Pinus taeda component was well constrained (within 1% of ground-based measurements). Although the closure of the C budget for the broadleaf species was poorer (within 20%), these species are a minor component of the forest. Under elevated CO₂, the C used annually for growth, turnover, and respiration balanced only 80% of the A_nC. Of the extra 700 g C m⁻² a⁻¹ (1999 and 2000 average), 86% is attributable to surface soil CO₂ efflux. This suggests that the production and turnover of fine roots was underestimated or that mycorrhizae and rhizodeposition became an increasingly important component of the C balance. Under elevated CO₂, net ecosystem production increased by 272 g C m⁻² a⁻¹: 44% greater than under ambient CO₂. The majority (87%) of this C was sequestered in a moderately long-term C pool in wood, with the remainder in the forest floor–soil subsystem.