Carbon dioxide fluxes in a spatially and temporally heterogeneous temperate grassland

Landscape position, grazing, and seasonal variation in precipitation and temperature create spatial and temporal variability in soil processes, and plant biomass and composition in grasslands. However, it is unclear how this variation in plant and soil properties affects carbon dioxide (CO₂) fluxes. The aim of this study is to explore the effect of grazing, topographic position, and seasonal variation in soil moisture and temperature on plant assimilation, shoot and soil respiration, and net ecosystem CO₂ exchange (NEE). Carbon dioxide fluxes, vegetation, and environmental variables were measured once a month inside and outside long-term ungulate exclosures in hilltop (dry) to slope bottom (mesic) grassland throughout the 2004 growing season in Yellowstone National Park. There was no difference in vegetation properties and CO₂ fluxes between the grazed and the ungrazed sites. The spatial and temporal variability in CO₂ fluxes were related to differences in aboveground biomass and total shoot nitrogen content, which were both related to variability in soil moisture. All sites were CO₂ sinks (NEE>0) for all our measurments taken throughout the growing season; but CO₂ fluxes were four- to fivefold higher at sites supporting the most aboveground biomass located at slope bottoms, compared to the sites with low biomass located at hilltops or slopes. The dry sites assimilated more CO₂ per gram aboveground biomass and stored proportionally more of the gross-assimilated CO₂ in the soil, compared to wet sites. These results indicate large spatio-temporal variability of CO₂ fluxes and suggest factors that control the variability in Yellowstone National Park.     

The use of soil respiration as an ecological indicator in arid ecosystems of the SE of Spain: Spatial variability and controlling factors

In the context of an ongoing monitoring study of the Cabo de Gata-Níjar Natural Park (SE of Spain), we explored the use of soil respiration as an indicator of ecosystem functioning reflecting changes in ecological processes in semiarid environments. With this purpose, we measured soil CO₂ efflux in six different and representative ecosystems of the Natural Park, with different land uses (forest and agricultural sites) and under different soil covers (under plant and bare soil) in two distinctive periods of the year: summer (dry period) and spring (growing season). We also measured the main soil properties and environmental variables. Soil CO₂ efflux ranged from 0.40 μmol m⁻² s⁻¹ in the dry period to 1.93 μmol m⁻² s⁻¹ in the growing season. Soil CO₂ efflux showed a large spatial variability, with different behaviour between the measured periods. Whereas in the dry period differences among ecosystems were larger (CVs 75-80%) than within them (CVs 40-55%), in the growing season the CVs were smaller (40-50%) and no differences were observed between or within ecosystem. The factors controlling soil CO₂ efflux also differed in the two measurement occasions. Whereas in the dry period soil CO₂ efflux was mainly the result of transport processes in the soil and therefore related to local factors (OC content, CN ratio, clay, rock outcrop, etc.) assigned to ecosystem conditions, in the growing season soil CO₂ efflux was dominated by soil CO₂ production and thus related only to organic carbon content and plant cover. In the growing season environmental variables explained ca. 10% of the variation in soil CO₂ efflux. In order to capture these different processes in different times of the year, i.e., diffusion versus production processes we calculated a new index, normalised seasonal difference in soil respiration (SDSR), which is proposed as a good indicator of the state and functioning of the ecosystem.     

Seasonal patterns in soil surface CO<Subscript>2</Subscript> flux under snow cover in 50 and 300 year old subalpine forests

Soil CO₂ flux can contribute as much as 60–80% of total ecosystem respiration in forests. Although considerable research has focused on quantifying this flux during the growing season, comparatively little effort has focused on non-growing season fluxes. We measured soil CO₂ efflux through snow in 50 and ~300 year old subalpine forest stands near Fraser CO. Our objectives were to quantify seasonal patterns in wintertime soil CO₂ flux; determine if differences in soil CO₂ flux between the two forest ages during the growing season persist during winter; and to quantify the sample size necessary to discern treatment differences. Soil CO₂ flux during the 2002–2003 and 2003–2004 snow season averaged 0.31 and 0.35 μmols m⁻² s⁻¹ for the young and old forests respectively; similar to the relative difference observed during summer. There was a significant seasonal pattern of soil CO₂ flux during the winter with fluxes averaging 0.22 μmols m⁻² s⁻¹ in December and January and increasing to an average of 0.61 μmols m⁻² s⁻¹ in May. Within-plot variability for measurements used in calculating flux was low. The coefficients of variation (CV) for CO₂ concentration, snowpack density, and snow depth were 17, 8 and 14%, respectively, yielding a CV for flux measurements within-plot of 29%. A within plot CV of 29% requires 8 sub-samples per plot to estimate the mean flux with a standard error of ±10% of the mean. Variability in CO₂ flux estimates among plots (size = 400 m²) was similar to that within plot and was also low (CV = ~28%). With a CV of 28% among plots, ten plots per treatment would have a 50% probability of detecting a 25% difference in treatment means for α = 0.05.     

Responses of Soil CO2 Fluxes to Short-Term Experimental Warming in Alpine Steppe Ecosystem,Northern Tibet

Soil carbon dioxide (CO₂) emission is one of the largest fluxes in the global carbon cycle. Therefore small changes in the size of this flux can have a large effect on atmospheric CO₂ concentrations and potentially constitute a powerful positive feedback to the climate system. Soil CO₂ fluxes in the alpine steppe ecosystem of Northern Tibet and their responses to short-term experimental warming were investigated during the growing season in 2011. The results showed that the total soil CO₂ emission fluxes during the entire growing season were 55.82 and 104.31 g C m^-2 for the control and warming plots, respectively. Thus, the soil CO₂ emission fluxes increased 86.86% with the air temperature increasing 3.74°C. Moreover, the temperature sensitivity coefficient (Q ₁₀) of the control and warming plots were 2.10 and 1.41, respectively. The soil temperature and soil moisture could partially explain the temporal variations of soil CO₂ fluxes. The relationship between the temporal variation of soil CO₂ fluxes and the soil temperature can be described by exponential equation. These results suggest that warming significantly promoted soil CO₂ emission in the alpine steppe ecosystem of Northern Tibet and indicate that this alpine ecosystem is very vulnerable to climate change. In addition, soil temperature and soil moisture are the key factors that controls soil organic matter decomposition and soil CO₂ emission, but temperature sensitivity significantly decreases due to the rise in temperature.     

Moisture Effects on Temperature Sensitivity of CO<Subscript>2</Subscript> Exchange in a Subarctic Heath Ecosystem

Carbon fluxes between natural ecosystems and the atmosphere have received increased attention in recent years due to the impact they have on climate. In order to investigate independently how soil moisture and temperature control carbon fluxes into and out of a dry subarctic dwarf shrub dominated heath, monoliths containing soil and plants were incubated at three different moisture levels and subjected to four different temperature levels between 7 and 20 °C. Ecosystem CO₂ exchange was monitored continuously day and night during the 16 to 18 days that each of three experiments lasted. Additionally, the carbon allocation pattern of the plants was investigated by labelling monoliths with ¹⁴CO₂ followed by harvest of above and below ground plant parts. The results revealed that the three different soil moisture levels caused distinctly differing levels of CO₂ fluxes. Also, both carbon fixation calculated as gross ecosystem production (GEP) and carbon release measured as ecosystem respiration (ER) increased with increasing temperatures, with ER increasing faster than GEP. Hence, short term carbon loss from the ecosystem accelerated with raised temperatures. Temperature sensitivity of the ecosystem was dependent on the soil moisture level, shown by differing Q10 values of both GEP and ER at different soil moisture levels. It is therefore highly important to take soil moisture levels into consideration when evaluating responses of ecosystem carbon balance to changes in temperature. The greatest C fixation took place via the two most dominant species of the ecosystem, Vaccinium uliginosum and Empetrum hermaphroditum, with the former being responsible for the different size of C fixation at the three moisture levels.     

Precipitation pulse size effects on Sonoran Desert soil microbial crusts

Deserts are characterized by low productivity and substantial unvegetated space, which is often covered by soil microbial crust communities. Microbial crusts are important for nitrogen fixation, soil stabilization and water infiltration, but their role in ecosystem production is not well understood. This study addresses the following questions: what are the CO₂ exchange responses of crusts to pulses of water, does the contribution of crusts to ecosystem flux differ from the soil respiratory flux, and is this contribution pulse size dependent? Following water application to crusts and soils, CO₂ exchange was measured and respiration was partitioned through mixing model analysis of Keeling plots across treatments. Following small precipitation pulse sizes, crusts contributed 80% of soil-level CO₂ fluxes to the atmosphere. However, following a large pulse event, roots and soil microbes contributed nearly 100% of the soil-level flux. Rainfall events in southern Arizona are dominated by small pulse sizes, suggesting that crusts may frequently contribute to ecosystem production. Carbon cycle studies of arid land systems should consider crusts as important contributors because of their dynamic responses to different pulse sizes as compared to the remaining ecosystem components.     

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.     

Ecosystem water fluxes for two grasslands in elevated CO2: a modeling analysis

The need to combine data from CO₂ field experiments with climate data remains urgent, particularly because each CO₂ experiment cannot run for decades to centuries. Furthermore, predictions for a given biome need to take into account differences in productivity and leaf area index (LAI) independent of CO₂-derived changes. In this study, we use long-term weather records and field data from the Jasper Ridge CO₂ experiment in Palo Alto, California, to model the effects of CO₂ and climate variability on ecosystem water fluxes. The sandstone and serpentine grasslands at Jasper Ridge provide a range of primary productivity and LAI, with the sandstone as the more productive system. Modeled soil water availability agreed well with published observations of time-domain reflectometry in the CO₂ experiment. Simulated water fluxes based on 10-year weather data (January 1985–December 1994) showed that the sandstone grassland had a much greater proportion of water movement through plants than did the serpentine; transpiration accounted for approximately 30% of annual fluxes in the sandstone and only 10% in the serpentine. Although simulated physiological and biomass changes were similar in both grasslands, the consequences of elevated CO₂ were greater for the sandstone water budget. Elevated CO₂ increased soil drainage by 20% in the sandstone, despite an approximately one-fifth increase in plant biomass; in the serpentine, drainage increased by <10% and soil evaporation was unchanged for the same simulated biomass change. Phenological changes, simulated by a 15-day lengthening of the growing season, had minimal impacts on the water budget. Annual variation in the timing and amount of rainfall was important for water fluxes in both grasslands. Elevated CO₂ increased sandstone drainage >50 mm in seven of ten years, but the relative increase in drainage varied from 10% to 300% depending on the year. Early-season transpiration in the sandstone decreased between 26% and 41%, with elevated CO₂ resulting in a simulated water savings of 54–76 mm. Even in years when precipitation was similar (e.g., 505 and 479 mm in years 3 and 4), the effect of CO₂ varied dramatically. The response of grassland water budgets to CO₂ depends on the productivity and structure of the grassland, the amount and timing of rainfall, and CO₂-induced changes in physiology. In systems with low LAI, large physiological changes may not necessarily alter total ecosystem water budgets dramatically. Received: 11 March 1997 / Accepted: 23 September 1997     

Alpine grassland ecosystem respiration variation under irrigation in Northern Tibet

Grassland ecosystems are important parts of terrestrial ecosystems and play an important role in the global carbon cycle. In recent years, the grasslands in Northern Tibet have experienced warming, and its precipitation has also increased. Alpine grassland irrigation measures could be a reasonable pathway to redistribute and make full use of the increased precipitation. In this study, we measured the soil respiration in alpine grassland in Northern Tibet under sprinkler head irrigation in the growing season to determine the relationships between soil temperature /water and ecosystem/soil respiration, soil moisture and Q₁₀, and soil temperature and Q₁₀. The results showed that after 2 years irrigation, alpine grassland aboveground biomass increased significantly, with 2010 higher than 2009. There was significant annual, seasonal and daily variation of soil respiration. Under irrigation, ecosystem respiration and soil respiration increased 75% and 64% respectively; soil water increase can promote the respiration of ecosystem and its components. In our results, the Q₁₀ value was 2.23–2.81, over the global average. The irrigation can promote ecosystem respiration temperature sensitivity. There was a positive linear correlation between ecosystem respiration and grassland aboveground biomass. The aboveground biomass accounted for 32.8% of ecosystem respiration variation. Soil respiration accounted for more than 70% of ecosystem respiration, indicating that the contribution to carbon emissions of soil respiration is very high. In short, we can project that in grasslands biomass and ecosystem respiration will increase under future precipitation change, which will significantly affect the function of alpine grassland carbon storage.     

Precipitation pulse use by an invasive woody legume: the role of soil texture and pulse size

Plant metabolic activity in arid and semi-arid environments is largely tied to episodic precipitation events or “pulses”. The ability of plants to take up and utilize rain pulses during the growing season in these water-limited ecosystems is determined in part by pulse timing, intensity and amount, and by hydrological properties of the soil that translate precipitation into plant-available soil moisture. We assessed the sensitivity of an invasive woody plant, velvet mesquite (Prosopis velutina Woot.), to large (35 mm) and small (10 mm) isotopically labeled irrigation pulses on two contrasting soil textures (sandy-loam vs. loamy-clay) in semi-desert grassland in southeastern Arizona, USA. Predawn leaf water potential (Ψ_pd), the isotopic abundance of deuterium in stem water (δD), the abundance of ¹³C in soluble leaf sugar (δ¹³C), and percent volumetric soil water content (θ_v) were measured prior to irrigation and repeatedly for 2 weeks following irrigation. Plant water potential and the percent of pulse water present in the stem xylem indicated that although mesquite trees on both coarse- and fine-textured soils quickly responded to the large irrigation pulse, the magnitude and duration of this response substantially differed between soil textures. After reaching a maximum 4 days after the irrigation, the fraction of pulse water in stem xylem decreased more rapidly on the loamy-clay soil than the sandy-loam soil. Similarly, on both soil textures mesquite significantly responded to the 10-mm pulse. However, the magnitude of this response was substantially greater for mesquite on the sandy-loam soil compared to loamy-clay soil. The relationship between Ψ_pd and δ¹³C of leaf-soluble carbohydrates over the pulse period did not differ between plants at the two sites, indicating that differences in photosynthetic response of mesquite trees to the moisture pulses was a function of soil water availability within the rooting zone rather than differences in plant biochemical or physiological constraints. Patterns of resource acquisition by mesquite during the dynamic wetting–drying cycle following rainfall pulses is controlled by a complex interaction between pulse size and soil hydraulic properties. A better understanding of how this interaction affects plant water availability and photosynthetic response is needed to predict how grassland structure and function will respond to climate change.     

Long‐term enhanced winter soil frost alters growing season CO2 fluxes through its impact on vegetation development in a boreal peatland

At high latitudes, winter climate change alters snow cover and, consequently, may cause a sustained change in soil frost dynamics. Altered winter soil conditions could influence the ecosystem exchange of carbon dioxide (CO₂) and, in turn, provide feedbacks to ongoing climate change. To investigate the mechanisms that modify the peatland CO₂ exchange in response to altered winter soil frost, we conducted a snow exclusion experiment to enhance winter soil frost and to evaluate its short‐term (1–3 years) and long‐term (11 years) effects on CO₂ fluxes during subsequent growing seasons in a boreal peatland. In the first 3 years after initiating the treatment, no significant effects were observed on either gross primary production (GPP) or ecosystem respiration (ER). However, after 11 years, the temperature sensitivity of ER was reduced in the treatment plots relative to the control, resulting in an overall lower ER in the former. Furthermore, early growing season GPP was also lower in the treatment plots than in the controls during periods with photosynthetic photon flux density (PPFD) ≥800 μmol m^?2 s^?1, corresponding to lower sedge leaf biomass in the treatment plots during the same period. During the peak growing season, a higher GPP was observed in the treatment plots under the low light condition (i.e. PPFD 400 μmol m^?2 s^?1) compared to the control. As Sphagnum moss maximizes photosynthesis at low light levels, this GPP difference between the plots may have been due to greater moss photosynthesis, as indicated by greater moss biomass production, in the treatment plots relative to the controls. Our study highlights the different responses to enhanced winter soil frost among plant functional types which regulate CO₂ fluxes, suggesting that winter climate change could considerably alter the growing season CO₂ exchange in boreal peatlands through its effect on vegetation development.     

Environmental effects on CO2 efflux from riparian tundra in the northern foothills of the Brooks Range,Alaska, USA

Summary Carbon dioxide efflux and soil microenvironmental factors were measured diurnally in Carex aquatilus-and Eriophorum angustifolium-dominated riparian tundra communities to determine the relative importance of soil environmental factors controlling ecosystem carbon dioxide exchange with the atmosphere. Measurements were made weekly between 18 June and 24 July 1990. Diurnal patterns in carbon dioxide efflux were best explained by changes in soil temperature, while seasonal changes in efflux were correlated with changes in depth to water table, depth to frozen soil and soil moisture. Carbon dioxide efflux rates were lowest early in the growing season when high water tables and low soil temperatures limited microbial and root activity. Individual rainfall events that raised the water table were found to strongly reduce carbon dioxide efflux. As the growing season progressed, rainfall was low and depth to water table and soil temperatures increased. In response, carbon dioxide efflux increased strongly, attaining rates late in the season of approximately 10 g CO₂ m^–2 day^–1. These rates are as high as maxima recorded for other arctic sites. A mathematical model is developed which demonstrates that soil temperature and depth to water table may be used as efficient predictors of ecosystem CO₂ efflux in this habitat. In parallel with the field measurements of CO₂ efflux, microbial respiration was studied in the laboratory as a function of temperature and water content. Estimates of microbial respiration per square meter under field conditions were made by adjusting for potential respiring soil volume as water table changed and using measured soil temperatures. The results indicate that the effect of these factors on microbial respiration may explain a large part of the diurnal and seasonal variation observed in CO₂ efflux. As in coastal tundra sites, environmental changes that alter water table depth in riparian tundra communities will have large effects on ecosystem CO₂ efflux and carbon balance.     

Effects of an increase in summer precipitation on leaf, soil, and ecosystem fluxes of CO2 and H2O in a sotol grassland in Big Bend National Park, Texas

Global climate models predict that in the next century precipitation in desert regions of the USA will increase, which is anticipated to affect biosphere/atmosphere exchanges of both CO₂ and H₂O. In a sotol grassland ecosystem in the Chihuahuan Desert at Big Bend National Park, we measured the response of leaf-level fluxes of CO₂ and H₂O 1 day before and up to 7 days after three supplemental precipitation pulses in the summer (June, July, and August 2004). In addition, the responses of leaf, soil, and ecosystem fluxes of CO₂ and H₂O to these precipitation pulses were also evaluated in September, 1 month after the final seasonal supplemental watering event. We found that plant carbon fixation responded positively to supplemental precipitation throughout the summer. Both shrubs and grasses in watered plots had increased rates of photosynthesis following pulses in June and July. In September, only grasses in watered plots had higher rates of photosynthesis than plants in the control plots. Soil respiration decreased in supplementally watered plots at the end of the summer. Due to these increased rates of photosynthesis in grasses and decreased rates of daytime soil respiration, watered ecosystems were a sink for carbon in September, assimilating on average 31 mmol CO₂ m⁻² s⁻¹ ground area day⁻¹. As a result of a 25% increase in summer precipitation, watered plots fixed eightfold more CO₂ during a 24-h period than control plots. In June and July, there were greater rates of transpiration for both grasses and shrubs in the watered plots. In September, similar rates of transpiration and soil water evaporation led to no observed treatment differences in ecosystem evapotranspiration, even though grasses transpired significantly more than shrubs. In summary, greater amounts of summer precipitation may lead to short-term increased carbon uptake by this sotol grassland ecosystem.     

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.     

The impact of invasion and subsequent removal of an exotic thistle, <Emphasis Type="Italic">Cynara cardunculus</Emphasis>, on CO<Subscript>2</Subscript> and H<Subscript>2</Subscript>O vapor exchange in a coastal California grassland

Changes in vegetation structure and composition, particularly due to the invasion of exotic species, are predicted to influence biosphere-atmosphere exchanges of mass and energy. Invasion of Cynara cardunculus (cardoon or artichoke thistle), a perennial, non-native thistle in coastal California grasslands presently dominated by non-native annual grasses, may alter rates of ecosystem CO₂ exchange and evapotranspiration (ET). During spring and summer 2006, we compared midday maximum net ecosystem CO₂ exchange (NEE) and ET among adjacent grassland plots where Cynara was present and where it was absent. Measurements of NEE supported the prediction that deeply-rooted Cynara increase midday ecosystem C-assimilation. Cynara-mediated shifts in NEE were associated with increases in ecosystem photosynthesis rather than changes in ecosystem respiration. Furthermore, the presence of Cynara was associated with increased ET during the growing season. An increase in aboveground live biomass (a proxy for leaf area) associated with Cynara invasion may underlie shifts in ecosystem CO₂ and water vapor exchange. Following mid-growing season sampling during April, we removed Cynara from half of the Cynara-containing plots with spot applications of herbicide. Three weeks later, midday fluxes in removal plots were indistinguishable from those in plots where Cynara was never present suggesting a lack of biogeochemical legacy effects. Similar to woody-encroachment in some semi-arid ecosystems, Cynara invasion increases midday ecosystem CO₂ assimilation and evapotranspiration rates and has the potential to increase C-storage in California coastal grasslands.     

The Effect of Increased N Deposition on Nitrous oxide, Methane and Carbon dioxide Fluxes from Unmanaged Forest and Grassland Communities in Michigan

Atmospheric nitrogen deposition is anticipated to increase over the next decades with possible implications for future forest-atmosphere interactions. Increased soil N₂O emissions, depressed CH₄ uptake and depressed soil respiration CO₂ loss is considered a likely response to increased N deposition. This study examined fluxes of N₂O, CH₄ and CO₂ over two growing seasons from soils in unmanaged forest and grassland communities on abandoned agricultural areas in Michigan. All sites were subject to simulated increased N-deposition in the range of 1–3 g N m⁻² annually. Nitrous oxide fluxes and soil N concentrations in coniferous and grassland sites were on the whole unaffected by the increased N-inputs. It is noteworthy though that N₂O emissions increased three-fold in the coniferous sites in the first growing season in response to the low N treatment, although the response was barely significant (p<0.06). In deciduous forests, we observed increased levels of soil mineral N during the second year of N fertilization, however N₂O fluxes did not increase. Rates of methane oxidation were similar in all sites with no affect of field N application. Likewise, we did not observe any changes in soil CO₂ efflux in response to N additions. The combination of tillage history and vegetation type was important for the trace gas fluxes, i.e. soil CO₂ efflux was greater in successional grassland sites compared with the forested sites and CH₄ uptake was reduced in post-tillage coniferous- and successional sites compared with the old-growth deciduous site. Our results indicate that short-term increased N availability influenced individual processes linked to trace gas turnover in the soil independently from the ecosystem N status. However, changes in whole system fluxes were not evident and were very likely mediated by competitive N uptake processes.     

Small-scale hydrological variation determines landscape CO2 fluxes in the high Arctic

We explored the influence of small-scale spatial variation in soil moisture on CO₂ fluxes in the high Arctic. Of five sites forming a hydrological gradient, CO₂ was emitted from the three driest sites and only the wettest site was a net sink of CO₂. Soil moisture was a good predictor of net ecosystem exchange (NEE). Higher gross ecosystem photosynthesis (GEP) was linked to higher bryophyte biomass and activity in response to the moisture conditions. Ecosystem respiration (R _e) rates increased with soil moisture until the soil became anaerobic and then R _e decreased. At well-drained sites R _e was driven by GEP, suggesting substrate and moisture limitation of soil respiration. We propose that spatial variability in soil moisture is a primary driver of NEE.     

Soil respiration response to three years of elevated CO2 and N fertilization in ponderosa pine (Pinus ponderosa Doug. ex Laws.)

We measured growing season soil CO₂ evolution under elevated atmospheric [CO₂] and soil nitrogen (N) additions. Our objectives were to determine treatment effects, quantify seasonal variation, and compare two measurement techniques. Elevated [CO₂] treatments were applied in open-top chambers containing ponderosa pine (Pinus ponderosa L.) seedlings. N applications were made annually in early spring. The experimental design was a replicated factorial combination of CO₂ (ambient, + 175, and +350 L L^–1 CO₂) and N (0, 10, and 20 g m^–2 N as ammonium sulphate). Soils were irrigated to maintain soil moisture at > 25 percent. Soil CO₂ evolution was measured over diurnal periods (20–22 hours) in October 1992, and April, June, and October 1993 and 1994 using a flow-through, infrared gas analyzer measurement system and corresponding pCO₂ measurements were made with gas wells. Significantly higher soil CO₂ evolution was observed in the elevated CO₂ treatments; N effects were not significant. Averaged across all measurement periods, fluxes, were 4.8, 8.0, and 6.5 for ambient + 175 CO₂, and +350 CO₂ respectively).Treatment variation was linearly related to fungal occurrence as observed in minirhizotron tubes. Seasonal variation in soil CO₂ evolution was non-linearly related to soil temperature; i.e., fluxes increased up to approximately soil temperature (10cm soil depth) and decreased dramatically at temperatures > 18°C. These patterns indicate exceeding optimal temperatures for biological activity. The dynamic, flow-through measurement system was weakly correlated (r = 0.57; p < 0.0001; n = 56) with the pCO₂ measurement method.     

Base Cation Cycling in a Pristine Watershed of the Canadian Boreal Forest

In forest ecosystems the single largest respiratory flux influencing net ecosystem productivity (NEP) is the total soil CO₂ efflux; however, it is difficult to make measurements of this flux that are accurate at the ecosystem scale. We examined patterns of soil CO₂ efflux using five different methods: auto-chambers, portable gas analyzers, eddy covariance along and two models parameterized with the observed data. The relation between soil temperature and soil moisture with soil CO₂ effluxes are also investigated, both inter-annually and seasonally, using these observations/results. Soil respiration rates (R _soil) are greatest during the growing season when soil temperatures are between 15 and 25 °C, but some soil CO₂ efflux occurs throughout the year. Measured soil respiration was sensitive to soil temperature, particularly during the spring and fall. All measurement methods produced similar annual estimates. Depending on the time of the year, the eddy covariance (flux tower) estimate for ecosystem respiration is similar to or slightly lower than estimates of annual soil CO₂ efflux from the other methods. As the eddy covariance estimate includes foliar and stem respiration which the other methods do not; it was expected to be larger (perhaps 15–30%). The auto-chamber system continuously measuring soil CO₂ efflux rates provides a level of temporal resolution that permits investigation of short- to longer term influences of factors on these efflux rates. The expense of building and maintaining an auto chamber system may not be necessary for those researchers interested in estimating R _soil annually, but auto-chambers do allow the capture of data from all seasons needed for model parameterization.     

Interactive Effects of Fire, Elevated Carbon Dioxide, Nitrogen Deposition, and precipitation on a California Annual Grassland

Although it is widely accepted that elevated atmospheric carbon dioxide (CO₂), nitrogen (N) deposition, and climate change will alter ecosystem productivity and function in the coming decades, the combined effects of these environmental changes may be nonadditive, and their interactions may be altered by disturbances, such as fire. We examined the influence of a summer wildfire on the interactive effects of elevated CO₂, N deposition, and increased precipitation in a full-factorial experiment conducted in a California annual grassland. In unburned plots, primary production was suppressed under elevated CO₂. Burning alone did not significantly affect production, but it increased total production in combination with nitrate additions and removed the suppressive effect of elevated CO₂. Increased production in response to nitrate in burned plots occurred as a result of the enhanced aboveground production of annual grasses and forbs, whereas the removal of the suppressive effect of elevated CO₂ occurred as a result of increased aboveground forb production in burned, CO₂-treated plots and decreased root production in burned plots under ambient CO₂.The tissue nitrogen–phosphorus ratio, which was assessed for annual grass shoots, decreased with burning and increased with nitrate addition. Burning removed surface litter from plots, resulting in an increase in maximum daily soil temperatures and a decrease in soil moisture both early and late in the growing season. Measures of vegetation greenness, based on canopy spectral reflectance, showed that plants in burned plots grew rapidly early in the season but senesced early. Overall, these results indicate that fire can alter the effects of elevated CO₂ and N addition on productivity in the short term, possibly by promoting increased phosphorus availability.