Species control variation in litter decomposition in a pine forest exposed to elevated CO2

Net primary production and the flux of dry matter and nutrients from vegetation to soils has increased following four years of exposure to elevated CO₂ in a southern pine forest in NC, USA. This has increased the demand for nutrients to support enhanced rates of NPP and altered the conditions for litter decomposition on the forest floor. We quantified the chemistry and decomposition dynamics of leaf litter produced by five of the most abundant tree species in this ecosystem during the third and fourth growing seasons under elevated CO₂. The objectives of this study were to determine (i) if there were systemic or species‐specific changes in leaf litter chemistry associated with a sustained enhancement of plant growth under elevated CO₂; and (ii) whether the process of litter decomposition was altered by increased inputs of energy and nutrients to the forest floor in the plots under elevated CO₂. Leaf litter chemistry, including various C fractions and N concentration, was virtually unchanged by elevated CO₂. With few exceptions, plant litter produced under elevated CO₂ lost mass or N at the same relative rate as that produced under ambient CO₂. The relationship between initial litter chemistry and decomposition was not altered by elevated CO₂. The greater forest floor mass and nutrient content in the plots under elevated CO₂ had no consistent or long‐term effect on litter decomposition. Thus, we found no evidence that plant and microbial processes under elevated CO₂ resulted in systemic changes in mass loss or N dynamics during decomposition. In contrast to the limited effects of elevated CO₂ on litter chemistry and decomposition, there were large differences among species in initial litter chemistry, mass loss and N dynamics during decomposition. If the species composition of this forest community is altered by elevated CO₂, the indirect effect of a change in species composition will exert greater control over the long‐term rate of nutrient cycling than the direct effect of elevated CO₂ on litter chemistry and decomposition dynamics alone.     

Decomposition of litter produced under elevated CO2: Dependence on plant species and nutrient supply

We investigated the effect of CO₂ concentration and soilnutrient availability during growth on the subsequent decomposition andnitrogen (N) release from litter of four annual grasses that differ inresource requirements and native habitat. Vulpia microstachys isa native grass found on California serpentine soils, whereas Avenafatua, Bromus hordaceus, and Lolium multiflorum areintroduced grasses restricted to more fertile sandstone soils (Hobbs & Mooney 1991). Growth in elevated CO₂ altered litter C:N ratio,decomposition, and N release, but the direction and magnitude of thechanges differed among plant species and nutrient treatments. ElevatedCO₂ had relatively modest effects on C:N ratio of litter,increasing this ratio in Lolium roots (and shoots at high nutrients),but decreasing C:N ratio in Avena shoots. Growth of plants underelevated CO₂ decreased the decomposition rate of Vulpialitter, but increased decomposition of Avena litter from the high-nutrient treatment. The impact of elevated CO₂ on N loss fromlitter also differed among species, with Vulpia litter from high-CO₂ plants releasing N more slowly than ambient-CO₂litter, whereas growth under elevated CO₂ caused increased Nloss from Avena litter. CO₂ effects on N release in Lolium and Bromus depended on the nutrient regime in whichplants were grown. There was no overall relationship between litter C:Nratio and decomposition rate or N release across species and treatments.Based on our study and the literature, we conclude that the effects ofelevated CO₂ on decomposition and N release from litter arehighly species-specific. These results do not support the hypothesis thatCO₂ effects on litter quality consistently lead to decreasednutrient availability in nutrient-limited ecosystems exposed to elevatedCO₂.     

The impact of elevated CO2, increased nitrogen availability and biodiversity on plant tissue quality and decomposition

Elevated CO₂, increased nitrogen (N) deposition and increasing species richness can increase net primary productivity (NPP). However, unless there are comparable changes in decomposition, increases in productivity will most likely be unsustainable. Without comparable increases in decomposition nutrients would accumulate in dead organic matter leading to nutrient limitations that could eventually prohibit additional increases in productivity. To address this issue, we measured aboveground plant and litter quality and belowground root quality, as well as decomposition of aboveground litter for one and 2‐year periods using in situ litterbags in response to a three‐way factorial manipulation of CO₂ (ambient vs. 560 ppm), N deposition (ambient vs. the addition of 4 g N m⁻² yr⁻¹) and plant species richness (one, four, nine and 16 species) in experimental grassland plots. Litter chemistry responded to the CO₂, N and plant diversity treatments, but decomposition was much less responsive. Elevated CO₂ induced decreases in % N and % lignin in plant tissues. N addition led to increases in % N and decreases in % lignin. Increasing plant diversity led to decreases in % N and % lignin and an increase in % cellulose. In contrast to the litter chemistry changes, elevated CO₂ had a much lower impact on decomposition and resulted in only a 2.5% decrease in carbon (C) loss. Detectable responses were not observed either to N addition or to species richness. These results suggest that global change factors such as biodiversity loss, elevated CO₂ and N deposition lead to significant changes in tissue quality; however, the response of decomposition is modest. Thus, the observed increases in productivity at higher diversity levels and with elevated CO₂ and N fertilization are not matched by an increase in decomposition rates. This lack of coupled responses between production and decomposition is likely to result in an accumulation of nutrients in the litter pool which will dampen the response of NPP to these factors over time.     

Diverse mechanisms for CO2 effects on grassland litter decomposition

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) can potentially alter litter decomposition rates by changing: (i) the litter quality of individual species, (ii) allocation patterns of individual species, (iii) the species composition of ecosystems (which could alter ecosystem‐level litter quality and allocation), (iv) patterns of soil moisture, and (v) the composition and size of microbial communities. To determine the relative importance of these mechanisms in a California annual grassland, we created four mixtures of litter that differed in species composition (the annual legume Lotus wrangelianus Fischer & C. Meyer comprised either 10% or 40% of the initial mass) and atmospheric [CO₂] during growth (ambient or double‐ambient). These mixtures decomposed for 33 weeks at three positions (above, on, and below the soil surface) in four types of grassland microcosms (fertilized and unfertilized microcosms exposed to elevated or ambient [CO₂]) and at a common field site. Initially, legume‐rich litter mixtures had higher nitrogen concentrations ([N]) than legume‐poor mixtures. In most positions and environments, the different litter mixtures decomposed at approximately the same rate. Fertilization and CO₂ enrichment of microcosms had no effect on mass loss of litter within them. However, mass loss was strongly related to litter position in both microcosms and the field. Nitrogen dynamics of litter were significantly related to the initial [N] of litter on the soil surface, but not in other positions. We conclude that changes in allocation patterns and species composition are likely to be the dominant mechanisms through which ecosystem‐level decomposition rates respond to increasing atmospheric [CO₂].     

Decomposition of secondary compounds from needle litter of Scots pine grown under elevated CO2 and O3

Elevated atmospheric carbon dioxide (CO₂) and ozone (O₃) concentrations have both been shown to affect plant tissue quality, which in turn could affect litter decomposition and carbon (C) and nutrient cycling. In order to evaluate effects of climate change on litter chemistry, needle litter was collected from Scots pine (Pinus sylvestris L.) saplings exposed to elevated CO₂ or O₃ concentration and their combination over three growing seasons in open‐top chambers. The decomposition of needle litter was followed for 19 months in a pine forest. During decomposition, needle samples for secondary compound analysis were collected and the mass loss of needles was followed. Main nutrients and total phenolics were analysed from litter in the beginning and at the end of the experiment. After 19‐month decomposition, the accumulated mass loss was about 34%; however, no significant differences were found in decomposition rates of needle litter between various treatments. Concentrations of total monoterpenes were about 4%, total resin acids 21% and total phenolics 14% of the initial concentrations in litter after 19‐month decomposition. In the beginning of litter decomposition, concentrations of individual monoterpenes –α‐pinene and β‐pinene – were significantly higher in needle litter grown under elevated CO₂. However, concentrations of total monoterpenes during the whole decomposition period were not significantly affected by CO₂ or O₃ treatments. Concentrations of some individual and total resin acids were higher in needle litter grown under elevated CO₂ or O₃ than under ambient air. There were no significant differences in concentrations of total phenolics as well as nitrogen (N) and the main nutrient concentrations between treatments during decomposition. High concentrations of monoterpenes and resin acids in needles might slightly delay C recycling in forest soils. It is concluded that elevated CO₂ and O₃ concentrations do not have remarkable impacts on litter decomposition processes in Scots pine forests.     

13C and 15N in microarthropods reveal little response of Douglas-fir ecosystems to climate change

Understanding ecosystem carbon (C) and nitrogen (N) cycling under global change requires experiments maintaining natural interactions among soil structure, soil communities, nutrient availability, and plant growth. In model Douglas-fir ecosystems maintained for five growing seasons, elevated temperature and carbon dioxide (CO₂) increased photosynthesis and increased C storage belowground but not aboveground. We hypothesized that interactions between N cycling and C fluxes through two main groups of microbes, mycorrhizal fungi (symbiotic with plants) and saprotrophic fungi (free-living), mediated ecosystem C storage. To quantify proportions of mycorrhizal and saprotrophic fungi, we measured stable isotopes in fungivorous microarthropods that efficiently censused the fungal community. Fungivorous microarthropods consumed on average 35% mycorrhizal fungi and 65% saprotrophic fungi. Elevated temperature decreased C flux through mycorrhizal fungi by 7%, whereas elevated CO₂ increased it by 4%. The dietary proportion of mycorrhizal fungi correlated across treatments with total plant biomass (n= 4, r²= 0.96, P= 0.021), but not with root biomass. This suggests that belowground allocation increased with increasing plant biomass, but that mycorrhizal fungi were stronger sinks for recent photosynthate than roots. Low N content of needles (0.8–1.1%) and A horizon soil (0.11%) coupled with high C : N ratios of A horizon soil (25–26) and litter (36–48) indicated severe N limitation. Elevated temperature treatments increased the saprotrophic decomposition of litter and lowered litter C : N ratios. Because of low N availability of this litter, its decomposition presumably increased N immobilization belowground, thereby restricting soil N availability for both mycorrhizal fungi and plant growth. Although increased photosynthesis with elevated CO₂ increased allocation of C to ectomycorrhizal fungi, it did not benefit plant N status. Most N for plants and soil storage was derived from litter decomposition. N sequestration by mycorrhizal fungi and limited N release during litter decomposition by saprotrophic fungi restricted N supply to plants, thereby constraining plant growth response to the different treatments.     

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.     

In situ litter decomposition and litter quality in a Mojave Desert ecosystem: effects of elevated atmospheric CO2 and interannual climate variability

Rising atmospheric CO₂ has been predicted to reduce litter decomposition as a result of CO₂‐induced reductions in litter quality. However, available data have not supported this hypothesis in mesic ecosystems, and no data are available for desert or semi‐arid ecosystems, which account for more than 35% of the Earth's land area. The objective of our study was to explore controls on litter decomposition in the Mojave Desert using elevated CO₂ and interannual climate variability as driving environmental factors. In particular, we sought to evaluate the extent to which decomposition is modulated by litter chemistry (C:N) and litter species and tissue composition. Naturally senesced litter was collected from each of nine 25 m diameter experimental plots, with six plots exposed to ambient [CO₂] or 367 μL CO₂ L^?1 and three plots continuously fumigated with elevated [CO₂] (550 μL CO₂ L^?1) using FACE technology beginning in April 1997. All litter collected in 1998 (a wet, or El Niño year; 306 mm precipitation) was pooled as was litter collected in 1999 (a dry year; 94 mm). Samples were allowed to decompose for 4 and 12 months starting in May 2001 in mesh litterbags in the locations from which litter was collected. Decomposition of litter produced under elevated CO₂ and ambient CO₂ did not differ. Litter produced in the wetter year showed more rapid initial decomposition (over the first 4 months) than that produced in the drier year (27±2% yr^?1 or 7.8±0.7 g m^?2 yr^?1 for 1998 litter; 18±3% yr^?1 or 2.2±0.4 g m^?2 yr^?1 for 1999 litter). C:N ratios of litter produced under elevated CO₂ (wet year: 37±0.5; dry year: 42±2.5) were higher than those of litter produced under ambient CO₂ (wet year: 34±1.1; dry year: 35±1.4). Litter production in the wet year (amb. CO₂: 25.1±1.1 g m^?2 yr^?1; elev. CO₂: 35.0±1.1 g m^?2 yr^?1) was more than twice as high as that in the dry year (amb. CO₂: 11.6±1.7 g m^?2, elev. CO₂: 13.3±3.4 g m^?2), and contained a greater proportion of Lycium pallidum and a lower proportion of Larrea tridentata than litter produced in the dry year. Decomposition, viewed across all treatments, decreased with increasing C:N ratios, decreased with increasing proportions of Larrea tridentata and increased with increasing proportions of Lycium pallidum and Lycium andersonii. Because litter C:N did not vary by litter production year, and CO₂ did not alter decomposition or litter species/tissue composition, it is likely that the impact of year‐to‐year variation in precipitation on the proportion of key plant species in the litter may be the most important way in which litter decomposition will be modulated in the Mojave Desert under future rising atmospheric CO₂.     

Elevated Carbon Dioxide and Litter Decomposition in California Annual Grasslands: Which Mechanisms Matter?

To date, most research that has examined the effect of elevated atmospheric carbon dioxide concentration ([CO₂]) on litter decomposition has focused on changes in the leaf litter quality of individual species. Results from California grasslands indicate that other CO₂ responses may have greater consequences for decomposition rates. For instance, CO₂-driven changes in either species dominance or patterns of biomass allocation would alter both the quality and the position of grassland litter. We review the results from studies in California grasslands to identify the mechanisms that affect grassland litter decomposition. We use a simple calculation that integrates the results of two studies to identify three mechanisms that have the potential to substantially alter decomposition rates as the atmospheric [CO₂] rises. Received 16 January 2001; accepted 26 September 2001.     

Effects of enhanced atmospheric CO2 and nutrient supply on the quality and subsequent decomposition of fine roots of Betula pendula Roth. and Picea sitchensis (Bong.) Carr.

Fine root litter derived from birch (Betula pendula Roth.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) plants grown under two CO₂ atmospheric concentrations (350 ppm and 600 ppm) and two nutrient regimes was used for decomposition studies in laboratory microcosms. Although there were interactions between litter type, CO₂/fertiliser treatments and decomposition rates, in general, an increase in the C/N ratio of the root tissue was observed for roots of both species grown under elevated CO₂ in unfertilized soil. Both weight loss and respiration of decomposing birch roots were significantly reduced in materials derived from enriched CO₂, whilst the decomposition of spruce roots showed no such effect. A parallel experiment was performed using Betula pendula root litter grown under different N regimes, in order to test the relationship between C/N ratio of litter and root decomposition rate. A highly significant (p<0.001) negative correlation between C/N ratio and root litter respiration was found, with an r²=0.97. The results suggest that the increased C/N ratio of plant tissues induced by elevated CO₂ can result in a reduction of decomposition rate, with a resulting increase in forest soil C stores.     

Microbial decomposition at elevated CO2 levels: effect of litter quality

The decomposition of senesced plant litter represents an important intermediate step in the cycling of nutrients between above- and below-ground systems. The rate of decomposition of plant litter is sensitive to fluctuations in a number of parameters, including environmental conditions, and particularly to changes in the quality of the litter. Increased C: N ratios of litter are thought to be one possible consequence of growth of plants under elevated [CO₂]. This response is likely to reduce the rate of decomposition of the litter. Evidence from the growth of plants in both pot and field studies suggests that growth of C3 plants in elevated atmospheric [CO₂] (600–700 μmol mol^–1) may lead to a significant increase in either/both the C: N and the lignin: N ratios of litter. Short-term decomposition of litter from plants showing this response in elevated [CO₂] has confirmed that decomposition occurs at a significantly lower rate. The limited studies of both the response of C4 plants to elevated [CO₂] and the subsequent degradability of the senescent litter suggest that no differences in litter quality or degradability occur. In terms of litter quality the response of plants therefore appears to be dependent upon photosynthetic type; the C:N and lignin:N ratios of litter from C3 plants exposed to elevated [CO₂] are increased, leading to lower degradation rates, while the nutrient ratios and degradation rates of litter from C4 plants grown in elevated [CO₂] remain unchanged. To date, very few ecosystem studies of decomposition have been carried out. Further work is required at the ecosystem level to determine whether the effects observed in laboratory, pot and field studies are also observed in long-term, complex ecosystem studies. Clearly if these results are repeated at the ecosystem level then significant changes in the cycling of C and N in important terrestrial ecosystems may occur as a results of elevated [CO₂].     

Elevated CO2 increases the long-term decomposition rate of Quercus myrtifolia leaf litter

Decomposition of Quercus myrtifolia leaf litter in a Florida scrub oak community was followed for 3 years in two separate experiments. In the first experiment, we examined the effects CO₂ and herbivore damage on litter quality and subsequent decomposition. Undamaged, chewed and mined litter generated under ambient and elevated (ambient+350 ppm V) CO₂ was allowed to decompose under ambient conditions for 3 years. Initial litter chemistry indicated that CO₂ levels had minor effects on litter quality. Litter damaged by leaf miners had higher initial concentrations of condensed tannins and nitrogen (N) and lower concentrations of hemicellulose and C : N ratios compared with undamaged and chewed litter. Despite variation in litter quality associated with CO₂, herbivory, and their interaction, there was no subsequent effect on rates of decomposition under ambient atmospheric conditions. In the second experiment, we examined the effects of source (ambient and elevated) of litter and decomposition site (ambient and elevated) on litter decomposition and N dynamics. Litter was not separated by damage type. The litter from both elevated and ambient CO₂ was then decomposed in both elevated and ambient CO₂ chambers. Initial litter chemistry indicated that concentrations of carbon (C), hemicellulose, and lignin were higher in litter from elevated than ambient CO₂ chambers. Despite differences in C and fiber concentrations, litter from ambient and elevated CO₂ decomposed at comparable rates. However, the atmosphere in which the decomposition took place resulted in significant differences in rates of decomposition. Litter decomposing under elevated CO₂ decomposed more rapidly than litter under ambient CO₂, and exhibited higher rates of mineral N accumulation. The results suggest that the atmospheric conditions during the decomposition process have a greater impact on rates of decomposition and N cycling than do the atmospheric conditions under which the foliage was produced.     

Litter decomposition rates in Canadian forests

The effect of litter quality and climate on the rate of decomposition of plant tissues was examined by the measurement of mass remaining after 3 years’ exposure of 11 litter types placed at 18 forest sites across Canada. Amongst sites, mass remaining was strongly related to mean annual temperature and precipitation and amongst litter types the ratio of Klason lignin to nitrogen in the initial tissue was the most important litter quality variable. When combined into a multiple regression, mean annual temperature, mean annual precipitation and Klason lignin:nitrogen ratio explained 73% of the variance in mass remaining for all sites and tissues. Using three doubled CO₂ GCM climate change scenarios for four Canadian regions, these relationships were used to predict increases in decomposition rate of 4–7% of contemporary rates (based on mass remaining after 3 years), because of increased temperature and precipitation. This increase may be partially offset by evidence that plants growing under elevated atmospheric CO₂ concentrations produce litter with high lignin:nitrogen ratios which slows the rate of decomposition, but this change will be small compared to the increased rate of decomposition derived from climatic changes.     

Seeing the forest for the trees: long-term exposure to elevated CO2 increases some herbivore densities

The effects of elevated CO₂ on plant growth and insect herbivory have been frequently investigated over the past 20 years. Most studies have shown an increase in plant growth, a decrease in plant nitrogen concentration, an increase in plant secondary metabolites and a decrease in herbivory. However, such studies have generally overlooked the fact that increases in plant production could cause increases of herbivores per unit area of habitat. Our study investigated leaf production, herbivory levels and herbivore abundance per unit area of leaf litter in a scrub‐oak system at Kennedy Space Center, Florida, under conditions of ambient and elevated CO₂, over an 11‐year period, from 1996 to 2007. In every year, herbivory, that is leafminer and leaftier abundance per 200 leaves, was lower under elevated CO₂ than ambient CO₂ for each of three species of oaks, Quercus myrtifolia, Quercus chapmanii and Quercus geminata. However, leaf litter production per 0.1143 m² was greater under elevated CO₂ than ambient CO₂ for Q. myrtifolia and Q. chapmanii, and this difference increased over the 11 years of the study. Leaf production of Q. geminata under elevated CO₂ did not increase. Leafminer densities per 0.1143 m² of litterfall for Q. myrtifolia and Q. chapmanii were initially lower under elevated CO₂. However, shortly after canopy closure in 2001, leafminer densities per 0.1143 m² of litter fall became higher under elevated CO₂ and remained higher for the remainder of the experiment. Leaftier densities per 0.1143 m² were also higher under elevated CO₂ for Q. myrtifolia and Q. chapmanii over the last 6 years of the experiment. There were no differences in leafminer or leaftier densities per 0.1143 m² of litter for Q. geminata. These results show three phenomena. First, they show that elevated CO₂ decreases herbivory on all oak species in the Florida scrub‐oak system. Second, despite lower numbers of herbivores per 200 leaves in elevated CO₂, increased leaf production resulted in higher herbivore densities per unit area of leaf litter for two oak species. Third, they corroborate other studies which suggest that the effects of elevated CO₂ on herbivores are species specific, meaning they depend on the particular plant species involved. Two oak species showed increases in leaf production and herbivore densities per 0.1143 m² in elevated CO₂ over time while another oak species did not. Our results point to a future world of elevated CO₂ where, despite lower plant herbivory, some insect herbivores may become more common.     

Short-term decomposition of litter produced by plants grown in ambient and elevated atmospheric CO2 concentrations

The effects of elevated atmospheric CO₂ (ambient + 340 μmol mol^–1) on above-ground litter decomposition were investigated over a 6-week period using a field-based mesocosm system. Soil respiratory activity in mesocosms incubated in ambient and elevated atmospheric CO₂ concentrations were not significantly different (t-test, P > 0.05) indicating that there were no direct effects of elevated atmospheric CO₂ on litter decomposition. A study of the indirect effects of CO₂ on soil respiration showed that soil mesocosms to which naturally senescent plant litter had been added (0.5% w/w) from the C₃ sedge Scirpus olneyi grown in elevated atmospheric CO₂ was reduced by an average of 17% throughout the study when compared to soil mesocosms to which litter from Scirpus olneyi grown in ambient conditions had been added. In contrast, similar experiments using senescent material from the C₄ grass Spartina patens showed no difference in soil respiration rates between mesocosms to which litter from plants grown in elevated or ambient CO₂ conditions had been added. Analysis of the C:N ratio and lignin content of the senescent material showed that, while the C:N ratio and lignin content of the Spartina patens litter did not vary with atmospheric CO₂ conditions, the C:N ratio (but not the lignin content) of the litter from Scirpus olneyi was significantly greater (t-test;P < 0.05) when derived from plants grown under elevated CO₂ (105:1 compared to 86:1 for litter derived from Scirpus olneyi grown under ambient conditions). The results suggest that the increased C:N ratio of the litter from the C₃ plant Scirpus olneyi grown under elevated CO₂ led to the lower rates of biodegradation observed as reduced soil respiration in the mesocosms. Further long-term experiments are now required to determine the effects of elevated CO₂ on C partitioning in terrestrial ecosystems.     

Impacts of elevated atmospheric CO2 on litter quality, litter decomposability and nitrogen turnover rate of two oak species in a Mediterranean forest ecosystem

Elevated CO₂ may affect litter quality of plants, and subsequently C and N cycling in terrestrial ecosystems, but changes in litter quality associated with elevated CO₂ are poorly known. Abscised leaf litter of two oak species (Quercus cerris L. and Q. pubescens Willd.) exposed to long-term elevated CO₂ around a natural CO₂ spring in Tuscany (Italy) was used to study the impact of increasing concentration of atmospheric CO₂ on litter quality and C and N turnover rates in a Mediterranean-type ecosystem. Litter samples were collected in an area with elevated CO₂ (>500 ppm) and in an area with ambient CO₂ concentration (360 ppm). Leaf samples were analysed for concentrations of total C, N, lignin, cellulose, acid detergent residue (ADR) and polyphenol. The decomposition rate of litter was studied using a litter bag experiment (12 months) and laboratory incubations (3 months). In the laboratory incubations, N mineralization in litter samples was measured as well (125 days). Litter quality was expressed in terms of chemical composition and element ratios. None of the litter quality parameters was affected by elevated CO₂ for the two Quercus species. Remaining mass in Q. cerris and Q. pubescens litter from elevated CO₂ was similar to that from ambient conditions. C mineralization in Q. pubescens litter from elevated CO₂ was lower than that from ambient CO₂, but the difference was insignificant. This effect was not observed for Q. cerris. N mineralization was higher from litter grown at elevated CO₂, but this difference disappeared at the end of the incubation. Litter of Q. pubescens had a higher quality than Q. cerris, and indeed mineralized more rapidly in the laboratory, but not under field conditions.     

Nitrogen cycling in grazed pastures at elevated CO2: N returns by ruminants

In pastures grazed by large herbivores, nutrients cycle both through litter and animal excreta. We compared nitrogen (N) returns from sheep grazing a temperate pasture exposed to ambient or elevated CO₂ (475 μmol mol^?1) in a FACE (Free Air CO₂ Enrichment) experiment established in the spring of 1997. In the spring of 2000 and 2001, we measured the chemical composition of the diet, sheep faeces and of individual plant species before grazing to characterize feed intake and to compare the intake of N to the N produced in faeces. In both years under elevated CO₂, leaves of the individual species exhibited lower N concentrations and higher water‐soluble carbohydrate (WSC) concentrations. There was a significantly greater proportion of legume in the diet at elevated CO₂ but, together with the changes in chemical composition of individual species, this resulted in diets that had similar N but higher WSC and digestibility for both ambient and elevated CO₂. We found that a greater proportion of dietary N was partitioned to urine at elevated CO₂, probably because of the higher proportion of legume N in the diet, with possible differences in protein quality. A potentially significant consequence of this change in partitioning is greater N loss through volatilization at higher CO₂ levels.     

Elevated [CO2], temperature increase and N supply effects on the turnover of below-ground carbon in a temperate grassland ecosystem

The effects of elevated [CO₂] (700 μl l^-1 CO₂) and temperature increase (+3 °C) on carbon turnover in grassland soils were studied during 2.5 years at two N fertiliser supplies (160 and 530 kg N ha^-1 y^-1) in an experiment with well-established ryegrass swards (Lolium perenne) supplied with the same amounts of irrigation water. During the growing season, swards from the control climate (350 μl l^-1 [CO₂] at outdoor air temperature) were pulse labelled by the addition of ¹³CO₂. The elevated [CO₂] treatments were continuously labelled by the addition of fossil-fuel derived CO₂ (^{13 C} of -40 to -50 ‰). Prior to the start of the experimental treatments, the carbon accumulated in the plant parts and in the soil macro-organic matter (‘old’ C) was at −32‰. During the experiment, the carbon fixed in the plant material (‘new’ C) was at −14 and −54‰ in the ambient and elevated [CO₂] treatments, respectively. During the experiment, the ¹³C isotopic mass balance method was used to calculate, for the top soil (0–15 cm), the carbon turnover in the stubble and roots and in the soil macro-organic matter above 200 μ (MOM). Elevated [CO₂] stimulated the turnover of organic carbon in the roots and stubble and in the MOM at N+, but not at N−. At the high N supply, the mean replacement time of ‘old’ C by ‘new’ C declined in elevated, compared to ambient [CO₂], from 18 to 7 months for the roots and stubble and from 25 to 17 months for the MOM. This resulted from increased rates of ‘new’ C accumulation and of ‘old’ C decay. By contrast, at the low N supply, despite an increase in the rate of accumulation of ‘new’ C, the soil C pools did not turnover faster in elevated [CO₂], as the rate of ‘old’ C decomposition was reduced. A 3 °C temperature increase in elevated [CO₂] decreased the input of fresh C to the roots and stubble and enhanced significantly the exponential rate for the ‘old’ C decomposition in the roots and stubble. An increased fertiliser N supply reduced the carbon turnover in the roots and stubble and in the MOM, in ambient but not in elevated [CO₂]. The respective roles for carbon turnover in the coarse soil OM fractions, of the C:N ratio of the litter, of the inorganic N availability and of a possible priming effect between C-substrates are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Drivers of increased soil respiration in a poplar coppice exposed to elevated CO2

Background and aims

The response of soil respiration (SR) to elevated CO₂ is driven by a number of processes and feedbacks. This work aims to i) detect the effect of elevated CO₂ on soil respiration during the second rotation of a short rotation forest, at two levels of N availability; and ii) identify the main drivers behind any changes in soil respiration.

Methods

A poplar plantation (POP-EUROFACE) was grown for two rotations of 3 years under elevated CO₂ maintained by a FACE (Free Air CO₂ Enrichment) technique. Root biomass, litter production and soil respiration were followed for two consecutive years after coppice.

Results

In the plantation, the stimulation of fine root and litter production under elevated CO₂ observed at the beginning of the rotation declined over time. Soil respiration (SR) was continuously stimulated by elevated CO₂, with a much larger enhancement during the growing (up to 111 %) than in the dormant season (40 %). The SR increase at first appeared to be due to the increase in fine root biomass, but at the end of the 2nd rotation was supported by litter decomposition and the availability of labile C. Soil respiration increase under elevated CO₂ was not affected by N availability.

Conclusions

The stimulation of SR by elevated CO₂ was sustained by the decomposition of above and belowground litter and by the greater availability of easily decomposable substrates into the soil. In the final year as elevated CO₂ did not increase C allocation to roots, the higher SR suggests greater C losses from the soil, thus reducing the potential for C accumulation.     

Elevated CO2 evokes quantitative and qualitative changes in carbon dynamics in a plant/soil system: mechanisms and implications

It is hypothesized that carbon storage in soil will increase under an elevated atmospheric CO₂ concentration due to a combination of an increased net CO₂ uptake, a shift in carbon allocation pattern in the plant/soil system and a decreased decomposition rate of plant residues. An overview of several studies, performed in our laboratory, on the effects of elevated CO₂ on net carbon uptake, allocation to the soil and decomposition of roots is given to test this hypothesis. The studies included wheat, ryegrass and Douglas-fir and comprised both short-term and long-term studies.Total dry weight of the plants increased up to 62%, but depended on nutrient availability. These results were supported by the data on net ¹⁴CO₂ uptake. A shift in ¹⁴C-carbon distribution from shoots to roots was found in perennial species, although this depended on nutrient availability.The decomposition experiments showed that roots cultivated at 700 L L^–1 CO₂ were decomposed more slowly than those cultivated at 350 L L^–1 CO₂. Even after two growing seasons differences up to 13% were observed, although this was found to be dependent on the nitrogen level at which the roots were grown.Both an increased carbon allocation to the soil due to an increased carbon uptake, whether or not combined with a shift in distribution pattern, and a decreased decomposition of root residues will enhance the possibilities of carbon sequestration in soil, thus supporting our hypothesis. However, nutrient availability and the response of the soil microbial biomass (size and activity) play a major role in the processes involved and require attention to clarify plant/soil responses in the long term with regard to sustained stimulation of carbon input into soils and the decomposability of roots and rhizodeposition. Soil texture will also have a strong effect on decomposition rates as a result of differences in the protecting capacity for organic matter. More detailed information on these changes is needed for a proper use of models simulating soil carbon dynamics in the long term.