Biotic degradation at night,abiotic degradation at day: positive feedbacks on litter decomposition in drylands

The arid and semi‐arid drylands of the world are increasingly recognized for their role in the terrestrial net carbon dioxide (CO₂) uptake, which depends largely on plant litter decomposition and the subsequent release of CO₂ back to the atmosphere. Observed decomposition rates in drylands are higher than predictions by biogeochemical models, which are traditionally based on microbial (biotic) degradation enabled by precipitation as the main mechanism of litter decomposition. Consequently, recent research in drylands has focused on abiotic mechanisms, mainly photochemical and thermal degradation, but they only partly explain litter decomposition under dry conditions, suggesting the operation of an additional mechanism. Here we show that in the absence of precipitation, absorption of dew and water vapor by litter in the field enables microbial degradation at night. By experimentally manipulating solar irradiance and nighttime air humidity, we estimated that most of the litter CO₂ efflux and decay occurring in the dry season was due to nighttime microbial degradation, with considerable additional contributions from photochemical and thermal degradation during the daytime. In a complementary study, at three sites across the Mediterranean Basin, litter CO₂ efflux was largely explained by litter moisture driving microbial degradation and ultraviolet radiation driving photodegradation. We further observed mutual enhancement of microbial activity and photodegradation at a daily scale. Identifying the interplay of decay mechanisms enhances our understanding of carbon turnover in drylands, which should improve the predictions of the long‐term trend of global carbon sequestration.     

Ecohydrological impacts of woody-plant encroachment: seasonal patterns of water and carbon dioxide exchange within a semiarid riparian environment

Across many dryland regions, historically grass‐dominated ecosystems have been encroached upon by woody‐plant species. In this paper, we compare ecosystem water and carbon dioxide (CO₂) fluxes over a grassland, a grassland–shrubland mosaic, and a fully developed woodland to evaluate potential consequences of woody‐plant encroachment on important ecosystem processes. All three sites were located in the riparian corridor of a river in the southwest US. As such, plants in these ecosystems may have access to moisture at the capillary fringe of the near‐surface water table. Using fluxes measured by eddy covariance in 2003 we found that ecosystem evapotranspiration (ET) and net ecosystem exchange of carbon dioxide (NEE) increased with increasing woody‐plant dominance. Growing season ET totals were 407, 450, and 639 mm in the grassland, shrubland, and woodland, respectively, and in excess of precipitation by 227, 265, and 473 mm. This excess was derived from groundwater, especially during the extremely dry premonsoon period when this was the only source of moisture available to plants. Access to groundwater by the deep‐rooted woody plants apparently decouples ecosystem ET from gross ecosystem production (GEP) with respect to precipitation. Compared with grasses, the woody plants were better able to use the stable groundwater source and had an increased net CO₂ gain during the dry periods. This enhanced plant activity resulted in substantial accumulation of leaf litter on the soil surface that, during rainy periods, may lead to high microbial respiration rates that offset these photosynthetic fluxes. March–December (primary growing season) totals of NEE were ?63, ?212, and ?233 g C m^?2 in the grassland, shrubland, and woodland, respectively. Thus, there was a greater disparity between ecosystem water use and the strength of the CO₂ sink as woody plants increased across the encroachment gradient. Despite a higher density of woody plants and a greater plant productivity in the woodland than in the shrubland, the woodland produced a larger respiration response to rainfall that largely offset its higher photosynthetic potential. These data suggest that the capacity for woody plants to exploit water resources in riparian areas results in enhanced carbon sequestration at the expense of increased groundwater use under current climate conditions, but the potential does not scale specifically as a function of woody‐plant abundance. These results highlight the important roles of water sources and ecosystem structure on the control of water and carbon balances in dryland areas.     

An accounting of C‐based trace gas release during abiotic plant litter degradation

Recent studies showed that photochemical breakdown (photodegradation) of plant material accounts for a substantial portion of litter decomposition and subsequent trace gas release in ecosystems under high radiative load and low precipitation. In the absence of solar radiation, thermal degradation may also cause trace gas release at temperatures below the ignition point. These observations suggest that the abiotic processes of photodegradation and thermal degradation of plant litter may be important in understanding global trace gas budgets. In a laboratory incubation study, we performed a simultaneous carbon (C) accounting of CO₂, CO, and CH₄ produced as a byproduct of photodegradation and thermal degradation of six different plant litter types that varied in chemical composition. The patterns of trace gas release during photodegradation and thermal degradation differed considerably across the six plant materials, suggesting that chemical composition of litter may influence the rates of abiotic degradation. There was a strong positive correlation between the rates of trace gas release during photodegradation and temperature. A significant portion of trace gases were produced during low temperature (< 100 °C) thermal degradation of litter in the absence of solar radiation, which was also positively correlated to temperature. In addition, both thermal degradation and photodegradation occurred in the absence of O₂. This indicates that the mechanism formerly accepted as photo‐oxidation may only be one of several photodegradation processes. We speculate that the direct breakdown of chemical groups such as carboxyl, carbonyl, and methoxyl groups may result in CO₂, CO, and CH₄ release. We suggest that the combined processes of thermal and photodegradation of litter may be a previously under accounted source of C‐based trace gases from terrestrial systems.     

Carbon limitation of soil respiration under winter snowpacks: potential feedbacks between growing season and winter carbon fluxes

A reduction in the length of the snow‐covered season in response to a warming of high‐latitude and high‐elevation ecosystems may increase soil carbon availability both through increased litter fall following longer growing seasons and by allowing early winter soil frosts that lyse plant and microbial cells. To evaluate how an increase in labile carbon during winter may affect ecosystem carbon balance we investigated the relationship between carbon availability and winter CO₂ fluxes at several locations in the Colorado Rockies. Landscape‐scale surveys of winter CO₂ fluxes from sites with different soil carbon content indicated that winter CO₂ fluxes were positively related to carbon availability and experimental additions of glucose to soil confirmed that CO₂ fluxes from snow‐covered soil at temperatures between 0 and ?3°C were carbon limited. Glucose added to snow‐covered soil increased CO₂ fluxes by 52–160% relative to control sites within 24 h and remained 62–70% higher after 30 days. Concurrently a shift in the δ¹³C values of emitted CO₂ toward the glucose value indicated preferential utilization of the added carbon confirming the presence of active heterotrophic respiration in soils at temperatures below 0°C. The sensitivity of these winter fluxes to substrate availability, coupled with predicted changes in winter snow cover, suggests that feedbacks between growing season carbon uptake and winter heterotrophic activity may have unforeseen consequences for carbon and nutrient cycling in northern forests. For example, published winter CO₂ fluxes indicate that on average 50% of growing season carbon uptake currently is respired during the winter; changes in winter CO₂ flux in response to climate change have the potential to reduce substantially the net carbon sink in these ecosystems.     

Carbon pools and fluxes along an environmental gradient in northern Arizona

Carbon pools and fluxes were quantified along an environmentalgradient in northern Arizona. Data are presented on vegetation, litter, andsoil C pools and soil CO₂ fluxesfrom ecosystems ranging from shrub-steppe through woodlands to coniferousforest and the ecotones in between. Carbon pool sizes and fluxes in thesesemiarid ecosystems vary with temperature and precipitation and are stronglyinfluenced by canopy cover. Ecosystem respiration is approximately 50percent greater in the more mesic, forest environment than in the dryshrub-steppe environment. Soil respiration rates within a site varyseasonally with temperature but appear to be constrained by low soilmoisture during dry summer months, when approximately 75% of totalannual soil respiration occurs. Total annual amount of CO₂ respired across all sites ispositively correlated with annual precipitation and negatively correlatedwith temperature. Results suggest that changes in the amount and periodicityof precipitation will have a greater effect on C pools and fluxes than willchanges in temperature in the semiarid Southwestern United States.     

Increased Quantity and Quality of Coarse Soil Organic Matter Fraction at Elevated CO2 in a Grazed Grassland are a Consequence of Enhanced Root Growth Rate and Turnover

The aims of this study were to determine whether elevated atmospheric CO₂ concentration modifies plant organic matter (OM) fluxes to the soil and whether any change in the fluxes can modify soil OM accumulation. Measurements were made in a grazed temperate grassland after almost 4 years exposure to elevated atmospheric CO₂ (475 μl l^-1) using a Free Air CO₂ Enrichment (FACE) facility located in the North Island of New Zealand. Aboveground herbage biomass and leaf litter production were not altered by elevated CO₂ but root growth rate, as measured with the ingrowth core method, and root turnover were strongly stimulated by elevated CO₂ particularly at low soil moisture contents during summer. Consequently, significantly more plant material was returned to the soil under elevated CO₂ leading to an accumulation of coarse (> 1 mm) particulate organic matter (POM) but not of finer POM fractions. The accumulating POM exhibited a lower C/N ratio, which was attributed to the higher proportion of legumes in the pasture under elevated CO₂. Only small changes were detected in the size and activity of the soil microbial biomass in response to the POM accumulation, suggesting that higher organic substrate availability did not stimulate microbial growth and activity despite the apparent lower C/N ratio of accumulating POM. As a result, elevated CO₂ may well lead to an accumulation of OM in grazed grassland soil in the long term.     

Annual burning of a tallgrass prairie inhibits C and N cycling in soil,increasing recalcitrant pyrogenic organic matter storage while reducing N availability

Grassland ecosystems store an estimated 30% of the world's total soil C and are frequently disturbed by wildfires or fire management. Aboveground litter decomposition is one of the main processes that form soil organic matter (SOM). However, during a fire biomass is removed or partially combusted and litter inputs to the soil are substituted with inputs of pyrogenic organic matter (py‐OM). Py‐OM accounts for a more recalcitrant plant input to SOM than fresh litter, and the historical frequency of burning may alter C and N retention of both fresh litter and py‐OM inputs to the soil. We compared the fate of these two forms of plant material by incubating ¹³C‐ and ¹⁵N‐labeled Andropogon gerardii litter and py‐OM at both an annually burned and an infrequently burned tallgrass prairie site for 11 months. We traced litter and py‐OM C and N into uncomplexed and organo‐mineral SOM fractions and CO₂ fluxes and determined how fire history affects the fate of these two forms of aboveground biomass. Evidence from CO₂ fluxes and SOM C:N ratios indicates that the litter was microbially transformed during decomposition while, besides an initial labile fraction, py‐OM added to SOM largely untransformed by soil microbes. Additionally, at the N‐limited annually burned site, litter N was tightly conserved. Together, these results demonstrate how, although py‐OM may contribute to C and N sequestration in the soil due to its resistance to microbial degradation, a long history of annual removal of fresh litter and input of py‐OM infers N limitation due to the inhibition of microbial decomposition of aboveground plant inputs to the soil. These results provide new insight into how fire may impact plant inputs to the soil, and the effects of py‐OM on SOM formation and ecosystem C and N cycling.     

Mind the gap: non‐biological processes contributing to soil CO2 efflux

Widespread recognition of the importance of soil CO₂ efflux as a major source of CO₂ to the atmosphere has led to active research. A large soil respiration database and recent reviews have compiled data, methods, and current challenges. This study highlights some deficiencies for a proper understanding of soil CO₂ efflux focusing on processes of soil CO₂ production and transport that have not received enough attention in the current soil respiration literature. It has mostly been assumed that soil CO₂ efflux is the result of biological processes (i.e. soil respiration), but recent studies demonstrate that pedochemical and geological processes, such as geothermal and volcanic CO₂ degassing, are potentially important in some areas. Besides the microbial decomposition of litter, solar radiation is responsible for photodegradation or photochemical degradation of litter. Diffusion is considered to be the main mechanism of CO₂ transport in the soil, but changes in atmospheric pressure and thermal convection may also be important mechanisms driving soil CO₂ efflux greater than diffusion under certain conditions. Lateral fluxes of carbon as dissolved organic and inorganic carbon occur and may cause an underestimation of soil CO₂ efflux. Traditionally soil CO₂ efflux has been measured with accumulation chambers assuming that the main transport mechanism is diffusion. New techniques are available such as improved automated chambers, CO₂ concentration profiles and isotopic techniques that may help to elucidate the sources of carbon from soils. We need to develop specific and standardized methods for different CO₂ sources to quantify this flux on a global scale. Biogeochemical models should include biological and non‐biological CO₂ production processes before we can predict the response of soil CO₂ efflux to climate change. Improving our understanding of the processes involved in soil CO₂ efflux should be a research priority given the importance of this flux in the global carbon budget.     

Trophic‐level dependent effects on CO2 emissions from experimental stream ecosystems

Concern over accelerating rates of species invasions and losses have initiated investigations into how local and global changes to predator abundance mediate trophic cascades that influence CO₂ fluxes of aquatic ecosystems. However, to date, no studies have investigated how species additions or losses at other consumer trophic levels influence the CO₂ flux of aquatic ecosystems. In this study, we added a large predatory stonefly, detritivorous stonefly, or grazer tadpole to experimental stream food webs and over a 70‐day period quantified their effects on community composition, leaf litter decomposition, chlorophyll‐a concentrations, and stream CO₂ emissions. In general, streams where the large grazer or large detritivore were added showed no change in total invertebrate biomass, leaf litter loss, chlorophyll‐a concentrations, or stream CO₂ emissions compared with controls; although we did observe a spike in CO₂ emissions in the large grazer treatment following a substantial reduction in chlorophyll‐a concentrations on day 28. However, the large grazer and large detritivore altered the community composition of streams by reducing the densities of other grazer and detritivore taxa, respectively, compared with controls. Conversely, the addition of the large predator created trophic cascades that reduced total invertebrate biomass and increased primary producer biomass. The cascading effects of the predator additions on the food web ultimately led to decreased CO₂ emissions from stream channels by up to 95%. Our results suggest that stream ecosystem processes were more influenced by changes in large predator abundance than large grazer or detritivore abundance, because of a lack of functionally similar large predators. Our study demonstrates that the presence/absence of species with unique functional roles may have consequences for the exchange of CO₂ between the ecosystem and the atmosphere.     

Increasing CO2 from subambient to elevated concentrations increases grassland respiration per unit of net carbon fixation

Respiration (carbon efflux) by terrestrial ecosystems is a major component of the global carbon (C) cycle, but the response of C efflux to atmospheric CO₂ enrichment remains uncertain. Respiration may respond directly to an increase in the availability of C substrates at high CO₂, but also may be affected indirectly by a CO₂‐mediated alteration in the amount by which respiration changes per unit of change in temperature or C uptake (sensitivity of respiration to temperature or C uptake). We measured CO₂ fluxes continuously during the final 2 years of a 4‐year experiment on C₃/C₄ grassland that was exposed to a 200–560 μmol mol^?1 CO₂ gradient. Flux measurements were used to determine whether CO₂ treatment affected nighttime respiration rates and the response of ecosystem respiration to seasonal changes in net C uptake and air temperature. Increasing CO₂ from subambient to elevated concentrations stimulated grassland respiration at night by increasing the net amount of C fixed during daylight and by increasing either the sensitivity of C efflux to daily changes in C fixation or the respiration rate in the absence of C uptake (basal ecosystem respiration rate). These latter two changes contributed to a 30–47% increase in the ratio of nighttime respiration to daytime net C influx as CO₂ increased from subamient to elevated concentrations. Daily changes in net C uptake were highly correlated with variation in temperature, meaning that the shared contribution of C uptake and temperature in explaining variance in respiration rates was large. Statistically controlling for collinearity between temperature and C uptake reduced the effect of a given change in C influx on respiration. Conversely, CO₂ treatment did not affect the response of grassland respiration to seasonal variation in temperature. Elevating CO₂ concentration increased grassland respiration rates by increasing both net C input and respiration per unit of C input. A better understanding of how C efflux varies with substrate supply thus may be required to accurately assess the C balance of terrestrial ecosystems.     

Sensitivity of organic matter decomposition to warming varies with its quality

The relationship between organic matter (OM) lability and temperature sensitivity is disputed, with recent observations suggesting that responses of relatively more resistant OM to increased temperature could be greater than, equivalent to, or less than responses of relatively more labile OM. This lack of clear understanding limits the ability to forecast carbon (C) cycle responses to temperature changes. Here, we derive a novel approach (denoted Q_10?q) that accounts for changes in OM quality during decomposition and use it to analyze data from three independent sources. Results from new laboratory soil incubations (labile Q_10?q=2.1 ± 0.2; more resistant Q_10?q=3.8 ± 0.3) and reanalysis of data from other soil incubations reported in the literature (labile Q_10?q=2.3; more resistant Q_10?q=3.3) demonstrate that temperature sensitivity of soil OM decomposition increases with decreasing soil OM lability. Analysis of data from a cross‐site, field litter bag decomposition study (labile Q_10?q=3.3 ± 0.2; resistant Q_10?q=4.9 ± 0.2) shows that litter OM follows the same pattern, with greater temperature sensitivity for more resistant litter OM. Furthermore, the initial response of cultivated soils, presumably containing less labile soil OM (Q_10?q=2.4 ± 0.3) was greater than that for undisturbed grassland soils (Q_10?q=1.7 ± 0.1). Soil C losses estimated using this approach will differ from previous estimates as a function of the magnitude of the temperature increase and the proportion of whole soil OM comprised of compounds sensitive to temperature over that temperature range. It is likely that increased temperature has already prompted release of significant amounts of C to the atmosphere as CO₂. Our results indicate that future losses of litter and soil C may be even greater than previously supposed.     

Litter Decomposition in a California Annual Grassland: Interactions Between Photodegradation and Litter Layer Thickness

In annual grasslands that experience a mediterranean-type climate, the synchrony between plant senescence and peak solar radiation over summer results in high litter sun exposure. We examined the decomposition of both shaded and sun-exposed litter over summer and inferred the effects of photodegradation from changes in mass loss and litter chemistry. The carry-over effects of summer litter exposure on wet season decomposition were also assessed, and the attenuation of photodegradation with litter layer thickness was used to estimate the proportion of grass litter lignin susceptible to photodegradation under different treatments of a factorial global change experiment. Over summer, mass loss from grass and forb litter exposed to ambient sunlight ranged from 8% to 10%, whereas lignin decreased in grass litter by approximately 20%. After one year of decomposition, mass losses from grass leaves exposed to sunlight over summer were more than double the mass losses from summer-shaded leaves. When shade litter layer thickness was varied, mass losses over summer for all treatments were also approximately 8%; however, lignin decreased significantly only in the low shade treatments (0–64 g m⁻² of shade litter). Aboveground production of annual grasses nearly quadrupled in response to the combined effects of N addition, elevated atmospheric CO₂, increased precipitation and warming. The estimated proportion of grass litter lignin experiencing full photodegradation ranged from 100% under ambient conditions to 31–62% in plots receiving the combined global change treatments. These results reveal an important role of sun exposure over summer in accelerating litter decomposition in these grasslands and provide evidence that future changes in the quantity of litter deposition may modulate the influence of photodegradation integrated across the litter layer.     

Landscape and Ecosystem-Level Controls on Net Carbon Dioxide Exchange along a Natural Moisture Gradient in Canadian Low Arctic Tundra

Our understanding of the controls and magnitudes of regional CO₂ exchanges in the Arctic are limited by uncertainties due to spatial heterogeneity in vegetation across the landscape and temporal variation in environmental conditions through the seasons. We measured daytime net ecosystem CO₂ exchange and each of its component fluxes in the three major tundra ecosystem-types that typically occur along natural moisture gradients in the Canadian Low Arctic biweekly during the full snow-free season of 2004. In addition, we used a plant-removal treatment to compare the contribution of bulk soil organic matter to total respiratory CO₂ loss among these ecosystems. Net CO₂ exchange rates varied strongly, but not consistently, among ecosystems in the spring and summer phases as a result of ecosystem-specific and differing responses of gross photosynthesis and respiration to temporal variation in environmental conditions. Overall, net carbon gain was largest in the wet sedge ecosystem and smallest in the dry heath. Our measures of CO₂ flux variation within each ecosystem were frequently most closely correlated with air or soil temperatures during each seasonal phase. Nevertheless, a particularly large rainfall event in early August rapidly decreased respiration rates and stimulated gross photosynthetic rates, resulting in peak rates of net carbon gain in all ecosystems. Finally, the bulk soil carbon contribution to total respiration was relatively high in the birch hummock ecosystem. Together, these results demonstrate that the relative influences of moisture and temperature as primary controls on daytime net ecosystem CO₂ exchange and its component fluxes differ in fundamental ways between the landscape and ecosystem scales. Furthermore, they strongly suggest that carbon cycling responses to environmental change are likely to be highly ecosystem-specific, and thus to vary substantially across the low arctic landscape. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

CO2 exchange in three Canadian High Arctic ecosystems: response to long-term experimental warming

Carbon dioxide exchange, soil C and N, leaf mineral nutrition and leaf carbon isotope discrimination (LCID‐Δ) were measured in three High Arctic tundra ecosystems over 2 years under ambient and long‐term (9 years) warmed (～2°C) conditions. These ecosystems are located at Alexandra Fiord (79°N) on Ellesmere Island, Nunavut, and span a soil water gradient; dry, mesic, and wet tundra. Growing season CO₂ fluxes (i.e., net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (R_e)) were measured using an infrared gas analyzer and winter C losses were estimated by chemical absorption. All three tundra ecosystems lost CO₂ to the atmosphere during the winter, ranging from 7 to 12 g CO₂‐C m^?2 season^?1 being highest in the wet tundra. The period during the growing season when mesic tundra switch from being a CO₂ source to a CO₂ sink was increased by 2 weeks because of warming and increases in GEP. Warming during the summer stimulated dry tundra GEP more than R_e and thus, NEE was consistently greater under warmed as opposed to ambient temperatures. In mesic tundra, warming stimulated GEP with no effect on R_e increasing NEE by ～10%, especially in the first half of the summer. During the ～70 days growing season (mid‐June–mid‐August), the dry and wet tundra ecosystems were net CO₂‐C sinks (30 and 67 g C m^?2 season^?1, respectively) and the mesic ecosystem was a net C source (58 g C m^?2 season^?1) to the atmosphere under ambient temperature conditions, due in part to unusual glacier melt water flooding that occurred in the mesic tundra. Experimental warming during the growing season increased net C uptake by ～12% in dry tundra, but reduced net C uptake by ～20% in wet tundra primarily because of greater rates of R_e as opposed to lower rates of GEP. Mesic tundra responded to long‐term warming with ～30% increase in GEP with almost no change in R_e reducing this tundra type to a slight C source (17 g C m^?2 season^?1). Warming caused LCID of Dryas integrafolia plants to be higher in dry tundra and lower in Salix arctic plants in mesic and wet tundra. Our findings indicate that: (1) High Arctic ecosystems, which occur in similar mesoclimates, have different net CO₂ exchange rates with the atmosphere; (2) long‐term warming can increase the net CO₂ exchange of High Arctic tundra by stimulating GEP, but it can also reduce net CO₂ exchange in some tundra types during the summer by stimulating R_e to a greater degree than stimulating GEP; (3) after 9 years of experimental warming, increases in soil carbon and nitrogen are detectable, in part, because of increases in deciduous shrub cover, biomass, and leaf litter inputs; (4) dry tundra increases in GEP, in response to long‐term warming, is reflected in D. integrifolia LCID; and (5) the differential carbon exchange responses of dry, mesic, and wet tundra to similar warming magnitudes appear to depend, in part, on the hydrologic (soil water) conditions. Annual net ecosystem CO₂‐C exchange rates ranged from losses of 64 g C m^?2 yr^?1 to gains of 55 g C m^?2 yr^?1. These magnitudes of positive NEE are close to the estimates of NPP for these tundra types in Alexandra Fiord and in other High Arctic locations based on destructive harvests.     

Diurnal and Seasonal Variations in Carbon Dioxide Exchange in Ecosystems in the Zhangye Oasis Area,Northwest China

Quantifying carbon dioxide exchange and understanding the response of key environmental factors in various ecosystems are critical to understanding regional carbon budgets and ecosystem behaviors. For this study, CO₂ fluxes were measured in a variety of ecosystems with an eddy covariance observation matrix between June 2012 and September 2012 in the Zhangye oasis area of Northwest China. The results show distinct diurnal variations in the CO₂ fluxes in vegetable field, orchard, wetland, and maize cropland. Diurnal variations of CO₂ fluxes were not obvious, and their values approached zero in the sandy desert, desert steppe, and Gobi ecosystems. Additionally, daily variations in the Gross Primary Production (GPP), Ecosystem Respiration (R_eco) and Net Ecosystem Exchange (NEE) were not obvious in the sandy desert, desert steppe, and Gobi ecosystems. In contrast, the distributions of the GPP, R_eco, and NEE show significant daily variations, that are closely related to the development of vegetation in the maize, wetland, orchard, and vegetable field ecosystems. All of the ecosystems are characterized by their carbon absorption during the observation period. The ability to absorb CO₂ differed significantly among the tested ecosystems. We also used the Michaelis-Menten equation and exponential curve fitting methods to analyze the impact of Photosynthetically Active Radiation (PAR) on the daytime CO₂ flux and impact of air temperature on R_eco at night. The results show that PAR is the dominant factor in controlling photosynthesis with limited solar radiation, and daytime CO₂ assimilation increases rapidly with PAR. Additionally, the carbon assimilation rate was found to increase slowly with high solar radiation. The light response parameters changed with each growth stage for all of the vegetation types, and higher light response values were observed during months or stages when the plants grew quickly. Light saturation points are different for different species. Nighttime R_eco increases exponentially with air temperature. High Q₁₀ values were observed when the vegetation coverage was relatively low, and low Q₁₀ values occurred when the vegetables grew vigorously.     

Due to symbiotic N2 fixation,five years of elevated atmospheric pCO2 had no effect on the N concentration of plant litter in fertile,mixed grassland

Experimental findings indicate that, in terrestrial ecosystems, nitrogen cycling changes under elevated partial pressure of atmospheric CO₂ (pCO₂). It was suggested that the concentration of N in plant litter as well as the amount of litter are responsible for these changes. However, for grassland ecosystems, there have been no relevant data available to support this hypothesis. Data from five years of the Swiss FACE experiment show that, under fertile soil conditions in a binary plant community consisting of Lolium perenne L. and Trifolium repens L., the concentration of litter N does not change under elevated atmospheric pCO₂; this applies to harvest losses, stubble, stolons and roots as the sources of litter. This is in strong contrast to the CO₂ response of L. perenne swards without associated legumes; in this case the above-ground concentration of biomass N decreased substantially. Increased symbiotic N₂ fixation in T. repens nodules and a greater proportion of the N-rich T. repens in the community are regarded as the main mechanisms that buffer the increased C introduction into the ecosystem under elevated atmospheric pCO₂. Our data also suggest that elevated atmospheric pCO₂ results in greater amounts of litter, mainly due to increased root biomass production. This study indicates that, in a fertile grassland ecosystem with legumes, the concentration of N in plant litter is not affected by elevated atmospheric pCO₂ and, thus, cannot explain CO₂-induced changes in the cycling of N. This revised version was published online in June 2006 with corrections to the Cover Date.     

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

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

The Role of Photodegradation in Surface Litter Decomposition Across a Grassland Ecosystem Precipitation Gradient

Differences in litter decomposition patterns among mesic, semiarid, and arid grassland ecosystems cannot be accurately explained by variation in temperature, moisture, and litter chemistry alone. We hypothesized that ultraviolet (UV) radiation enhances decomposition in grassland ecosystems via photodegradation, more so in arid compared to mesic ecosystems, and in litter that is more recalcitrant to microbial decomposition (with high compared to low lignin concentrations). In a 2-year field study, we manipulated the amount of UV radiation reaching the litter layer at three grassland sites in Minnesota, Colorado, and New Mexico, USA, that represented mesic, semiarid, and arid grassland ecosystems, respectively. Two common grass leaf litter types of contrasting lignin:N were placed at each site under screens that either passed all solar radiation wavelengths or passed all but UV wavelengths. Decomposition was generally faster when litter was exposed to UV radiation across all three sites. In contrast to our hypothesis, the contribution of photodegradation in the decomposition process was not consistently greater at the more arid sites or for litter with higher lignin content. Additionally, at the most arid site, exposure to UV radiation could not explain decomposition rates that were faster than expected given climate constraints or lack of N immobilization by decomposing litter. Although photodegradation plays an important role in the decomposition process in a wider range of grassland sites than previously documented, it does not fully explain the differences in decomposition rates among grassland ecosystems of contrasting aridity.     

Fluxes of carbon, water and energy over Brazilian cerrado: an analysis using eddy covariance and stable isotopes

We present the energy and mass balance of cerrado sensu stricto (a Brazilian form of savanna), in which a mixture of shrubs, trees and grasses forms a vegetation with a leaf area index of 1·0 in the wet season and 0·4 in the dry season. In the wet season the available energy was equally dissipated between sensible heat and evaporation, but in the dry season at high irradiance the sensible heat greatly exceeded evaporation. Ecosystem surface conductance g_s in the wet season rose abruptly to 0·3 mol m^?2 s^?1 and fell gradually as the day progressed. Much of the total variation in g_s was associated with variation in the leaf-to-air vapour pressure deficit of water and the solar irradiance. In the dry season the maximal g_s values were only 0·1 mol m^?2 s^?1. Maximal net ecosystem fluxes of CO₂ in the wet and dry season were –10 and –15 μmol CO₂ m^?2 s^?1, respectively (sign convention: negative denotes fluxes from atmosphere to vegetation). The canopy was well coupled to the atmosphere, and there was rarely a significant build-up of respiratory CO₂ during the night. For observations in the wet season, the vegetation was a carbon dioxide sink, of maximal strength 0·15 mol m^?2 d^?1. However, it was a source of carbon dioxide for a brief period at the height of the dry season. Leaf carbon isotopic composition showed all the grasses except for one species to be C₄, and all the palms and woody plants to be C₃. The CO₂ coming from the soil had an isotopic composition that suggested 40% of it was of C₄ origin.     

Spatial patterns of litter decomposition in the littoral zone of boreal lakes

1. We studied the patterns of litter decomposition in lake littoral habitats and investigated whether decay rates, as an integrating proxy for environmental conditions in the sediment, would co‐vary with net carbon dioxide (CO₂) exchange and methane (CH₄) efflux. These gas fluxes are known to be sensitive to environmental conditions. Losses in the mass of cellulose, root, rhizome and moss litter were measured during 2 years in boreal littoral wetlands in Finland and compared with published data on concurrently measured gas fluxes. Four study sites covered a range of sediment types and hydrological conditions. 2. Decomposition was not linearly related to the duration of flooding but depended on sediment type. Readily decomposable litter fractions, such as cellulose and rhizome litter, lost mass at a faster rate in marshes with a longer period of flooding but wide water level fluctuations that hinder establishment of a Sphagnum cover, than in peat‐forming fens. In marshes, the mean first‐year mass losses were 83–99% and 19–62% for cellulose and rhizomes, respectively. In fens, the respective losses were 40–53% and 33%. In the first year, the loss in the mass of the more recalcitrant root litter did not differ between sites (mean 19–30%) and moss litter lost no mass. 3. The estimated first‐year carbon loss from belowground litter was about 0.1–0.3 times ecosystem respiration and roughly similar to net carbon gas (CO₂, CH₄) efflux, suggesting that vascular plants and recent plant residues contribute substantially to ecosystem release of carbon gases. On the other hand, at least 40% of the mass of the belowground litter remained on a littoral site after the first 2 years of decomposition. Slow decomposition may indicate the accumulation of organic‐rich sediments. The accumulated carbon could explain the excess CO₂ release found in most littoral sites. In continuously inundated sites decomposition rates were similar to those in periodically flooded sites, but ecosystem‐atmosphere CO₂ exchange fell to close to zero. This discrepancy implies that the released CO₂ is dissolved in water and may be exported into the pelagic zone of the lake.