Deeper Snow Enhances Winter Respiration from Both Plant-associated and Bulk Soil Carbon Pools in Birch Hummock Tundra

It has only recently become apparent that biological activity during winter in seasonally snow-covered ecosystems may exert a significant influence on biogeochemical cycling and ecosystem function. One-seventh of the global soil carbon pool is stored in the bulk soil component of arctic ecosystems. Consistent climate change predictions of substantial increases in winter air temperatures and snow depths for the Arctic indicate that this region may become a significant net annual source of CO₂ to the atmosphere if its bulk soil carbon is decomposed. We used snow fences to investigate the influence of a moderate increase in snow depth from approximately 0.3 m (ambient) to approximately 1 m on winter carbon dioxide fluxes from mesic birch hummock tundra in northern Canada. We differentiated fluxes derived from the bulk soil and plant-associated carbon pools using an experimental ‘weeding’ manipulation. Increased snow depth enhanced the wintertime carbon flux from both pools, strongly suggesting that respiration from each was sensitive to warmer soil temperatures. Furthermore, deepened snow resulted in cooler and relatively stable soil temperatures during the spring-thaw period, as well as delayed and fewer freeze–thaw cycles. The snow fence treatment increased mean total winter efflux from 27 to 43 g CO₂-C m⁻². Because total 2004 growing season net ecosystem exchange for this site is estimated at 29–37 g CO₂-C m⁻², our results strongly suggest that a moderate increase in snow depth can enhance winter respiration sufficiently to switch the ecosystem annual net carbon exchange from a sink to source, resulting in net CO₂ release to the atmosphere.     

Impacts of Elevated Carbon Dioxide and Temperature on a Boreal Forest Ecosystem (CLIMEX Project)

To evaluate the effects of climate change on boreal forest ecosystems, both atmospheric CO₂ (to 560 ppmv) and air temperature (by 3°–5°C above ambient) were increased at a forested headwater catchment in southern Norway. The entire catchment (860 m²) is enclosed within a transparent greenhouse, and the upper 20% of the catchment area is partitioned such that it receives no climate treatment and serves as an untreated control. Both the control and treatment areas inside the greenhouse receive deacidified rain. Within 3 years, soil nitrogen (N) mineralization has increased and the growing season has been prolonged relative to the control area. This has helped to sustain an increase in plant growth relative to the control and has also promoted increased N export in stream water. Photosynthetic capacity and carbon–nitrogen ratio of new leaves of most plant species did not change. While the ecosystem now loses N, the long-term fate of soil N is a key uncertainty in predicting the future response of boreal ecosystems to climate change. Received 18 November 1997; accepted 13 April 1998.     

Temperature and substrate controls on intra-annual variation in ecosystem respiration in two subarctic vegetation types

Arctic ecosystems are important in the context of climate change because they are expected to undergo the most rapid temperature increases, and could provide a globally significant release of CO₂ to the atmosphere from their extensive bulk soil organic carbon reserves. Understanding the relative contributions of bulk soil organic matter and plant‐associated carbon pools to ecosystem respiration is critical to predicting the response of arctic ecosystem net carbon balance to climate change. In this study, we determined the variation in ecosystem respiration rates from birch forest understory and heath tundra vegetation types in northern Sweden through a full annual cycle. We used a plant biomass removal treatment to differentiate bulk soil organic matter respiration from total ecosystem respiration in each vegetation type. Plant‐associated and bulk soil organic matter carbon pools each contributed significantly to ecosystem respiration during most phases of winter and summer in the two vegetation types. Ecosystem respiration rates through the year did not differ significantly between vegetation types despite substantial differences in biomass pools, soil depth and temperature regime. Most (76–92%) of the intra‐annual variation in ecosystem respiration rates from these two common mesic subarctic ecosystems was explained using a first‐order exponential equation relating respiration to substrate chemical quality and soil temperature. Removal of plants and their current year's litter significantly reduced the sensitivity of ecosystem respiration to intra‐annual variations in soil temperature for both vegetation types, indicating that respiration derived from recent plant carbon fixation was more temperature sensitive than respiration from bulk soil organic matter carbon stores. Accurate assessment of the potential for positive feedbacks from high‐latitude ecosystems to CO₂‐induced climate change will require the development of ecosystem‐level physiological models of net carbon exchange that differentiate the responses of major C pools, that account for effects of vegetation type, and that integrate over summer and winter seasons.     

Plant Species Composition and Productivity following Permafrost Thaw and Thermokarst in Alaskan Tundra

Climate warming is expected to have a large impact on plant species composition and productivity in northern latitude ecosystems. Warming can affect vegetation communities directly through temperature effects on plant growth and indirectly through alteration of soil nutrient availability. In addition, warming can cause permafrost to thaw and thermokarst (ground subsidence) to develop, which can alter the structure of the ecosystem by altering hydrological patterns within a site. These multiple direct and indirect effects of permafrost thawing are difficult to simulate in experimental approaches that often manipulate only one or two factors. Here, we used a natural gradient approach with three sites to represent stages in the process of permafrost thawing and thermokarst. We found that vascular plant biomass shifted from graminoid-dominated tundra in the least disturbed site to shrub-dominated tundra at the oldest, most subsided site, whereas the intermediate site was co-dominated by both plant functional groups. Vascular plant productivity patterns followed the changes in biomass, whereas nonvascular moss productivity was especially important in the oldest, most subsided site. The coefficient of variation for soil moisture was higher in the oldest, most subsided site suggesting that in addition to more wet microsites, there were other microsites that were drier. Across all sites, graminoids preferred the cold, dry microsites whereas the moss and shrubs were associated with the warm, moist microsites. Total nitrogen contained in green plant biomass differed across sites, suggesting that there were increases in soil nitrogen availability where permafrost had thawed.     

Ecosystem CO2 production during winter in a Swedish subarctic region: the relative importance of climate and vegetation type

General circulation models consistently predict that regional warming will be most rapid in the Arctic, that this warming will be predominantly in the winter season, and that it will often be accompanied by increasing snowfall. Paradoxically, despite the strong cold season emphasis in these predictions, we know relatively little about the plot and landscape‐level controls on tundra biogeochemical cycling in wintertime as compared to summertime. We investigated the relative influence of vegetation type and climate on CO₂ production rates and total wintertime CO₂ release in the Scandinavian subarctic. Ecosystem respiration rates and a wide range of associated environmental and substrate pool size variables were measured in the two most common vegetation types of the region (birch understorey and heath tundra) at four paired sites along a 50 km transect through a strong snow depth gradient in northern Sweden. Both climate and vegetation type were strong interactive controls on ecosystem CO₂ production rates during winter. Of all variables tested, soil temperature explained by far the largest amount of variation in respiration rates (41–75%). Our results indicate that vegetation type only exerted an influence on respiration when snow depth was below a certain threshold (～1 m). Thus, tall vegetation that enhanced snow accumulation within that threshold resulted in more effective thermal insulation from severe air temperatures, thereby significantly increasing respiratory activity. At the end of winter, within several days of snowmelt, gross ecosystem photosynthesis rates were of a similar magnitude to ecosystem respiration, resulting in significant net carbon gain in some instances. Finally, climate and vegetation type were also strong interactive controls on total wintertime respiration, suggesting that spatial variations in maximum snowdepth may be a primary determinant of regional patterns of wintertime CO₂ release. Together, our results have important implications for predictions of how the distribution of tundra vegetation types and the carbon balances of arctic ecosystems will respond to climate change during winter because they indicate a threshold (～1 m) above which there would be little effect of increased snow accumulation on wintertime biogeochemical cycling.     

Carbon Dioxide Dynamics and Controls in a Deep-water Wetland on the Qinghai-Tibetan Plateau

To initially characterize the dynamics and environmental controls of CO₂, ecosystem CO₂ fluxes were measured for different vegetation zones in a deep-water wetland on the Qinghai-Tibetan Plateau during the growing season of 2002. Four zones of vegetation along a gradient from shallow to deep water were dominated, respectively by the emergent species Carex allivescens V. Krez., Scirpus distigmaticus L., Hippuris vulgaris L., and the submerged species Potamogeton pectinatus L. Gross primary production (GPP), ecosystem respiration (Re), and net ecosystem production (NEP) were markedly different among the vegetation zones, with lower Re and GPP in deeper water. NEP was highest in the Scirpus-dominated zone with moderate water depth, but lowest in the Potamogeton-zone that occupied approximately 75% of the total wetland area. Diurnal variation in CO₂ flux was highly correlated with variation in light intensity and soil temperature. The relationship between CO₂ flux and these environmental variables varied among the vegetation zones. Seasonal CO₂ fluxes, including GPP, Re, and NEP, were strongly correlated with aboveground biomass, which was in turn determined by water depth. In the early growing season, temperature sensitivity (Q₁₀) for Re varied from 6.0 to 8.9 depending on vegetation zone. Q₁₀ decreased in the late growing season. Estimated NEP for the whole deep-water wetland over the growing season was 24 g C m⁻². Our results suggest that water depth is the major environmental control of seasonal variation in CO₂ flux, whereas photosynthetic photon flux density (PPFD) controls diurnal dynamics.     

Primary Productivity and Water Balance of Grassland Vegetation on Three Soils in a Continuous CO2 Gradient: Initial Results from the Lysimeter CO2 Gradient Experiment

Field studies of atmospheric CO₂ effects on ecosystems usually include few levels of CO₂ and a single soil type, making it difficult to ascertain the shape of responses to increasing CO₂ or to generalize across soil types. The Lysimeter CO₂ Gradient (LYCOG) chambers were constructed to maintain a linear gradient of atmospheric CO₂ (~250 to 500 μl l⁻¹) on grassland vegetation established on intact soil monoliths from three soil series. The chambers maintained a linear daytime CO₂ gradient from 263 μl l⁻¹ at the subambient end of the gradient to 502 μl l⁻¹ at the superambient end, as well as a linear nighttime CO₂ gradient. Temperature variation within the chambers affected aboveground biomass and evapotranspiration, but the effects of temperature were small compared to the expected effects of CO₂. Aboveground biomass on Austin soils was 40% less than on Bastrop and Houston soils. Biomass differences between soils resulted from variation in biomass of Sorghastrum nutans, Bouteloua curtipendula, Schizachyrium scoparium (C₄ grasses), and Solidago canadensis (C₃ forb), suggesting the CO₂ sensitivity of these species may differ among soils. Evapotranspiration did not differ among the soils, but the CO₂ sensitivity of leaf-level photosynthesis and water use efficiency in S. canadensis was greater on Houston and Bastrop than on Austin soils, whereas the CO₂ sensitivity of soil CO₂ efflux was greater on Bastrop soils than on Austin or Houston soils. The effects of soil type on CO₂ sensitivity may be smaller for some processes that are tightly coupled to microclimate. LYCOG is useful for discerning the effects of soil type on the CO₂ sensitivity of ecosystem function in grasslands. Author Contributions: PF conceived study, analyzed data, and wrote the paper. AK, AP analyzed data. DH, VJ, RJ, HJ, and WP conceived study, and conducted research.     

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.     

青藏高原气候变化与高寒草地

综述了近五十年来青藏高原气候和高寒草地的变化趋势,阐述了气候变化对高寒草地的可能影响。气候变化主要通过水、热过程及其诱导的环境变化对青藏高原高寒草地产生显著的影响。主要过程包括：气候变化对气候带、植被带、植物、植物群落、农业生产以及生态系统固碳潜力等的影响。从目前的观测和研究结果来看,有关青藏高原气候变化及其对高寒草地的可能影响都还很难得出一致的结论。因此,如何科学评价气候变化及其预测和评价对高寒草地结构和功能的潜在影响,以及如何将已经发生的变化纳入到全球变化模型或评价体系中,以便更加精确地评估气候变化的长期影响,将成为必须要回答的关键科学问题。     

Concentration and Fluxes of Dissolved Organic Carbon (DOC) in Three Norway Spruce Stands along a Climatic Gradient in Sweden

Leaching of dissolved organic carbon (DOC) from the forest floor and transport in soil solution into the mineral soil are important for carbon cycling in boreal forest ecosystems. We examined DOC concentrations in bulk deposition, throughfall and in soil solutions collected under the O and B horizons in three Norway spruce stands along a climatic gradient in Sweden. Mean annual temperature for the three sites was 5.5, 3.4 and 1.2 °C. At each site we also examined the effect of soil moisture on DOC dynamics along a moisture gradient (dry, mesic and moist plots). To obtain information about the fate of DOC leached from the O horizon into the mineral soil, ¹⁴C measurements were made on bulk organic matter and DOC. The concentration and fluxes of DOC in O horizon leachates were highest at the southern site and lowest at the northern. Average DOC concentrations at the southern, central and northern sites were 49, 39 and 30 mg l⁻¹, respectively. We suggest that DOC leaching rates from O horizons were related to the net primary production of the ecosystem. Soil temperature probably governed the within-year variation in DOC concentration in O horizon leachates, but the peak in DOC was delayed relative to that of temperature, probably due to sorption processes. Neither soil moisture regime (dry, mesic or moist plots) nor seasonal variation in soil moisture seemed to be of any significance for the concentration of DOC leached from the O horizon. The ¹⁴C measurements showed that DOC in soil solution collected below the B horizon was derived mainly from the B horizon itself, rather than from the O horizon, indicating a substantial exchange (sorption–desorption reactions) between incoming DOC and soil organic carbon in the mineral soil.     

Diurnal and seasonal variation in methane emissions in a northern Canadian peatland measured by eddy covariance

Eddy covariance measurements of methane (CH₄) net flux were made in a boreal fen, typical of the most abundant peatlands in western Canada during May–September 2007. The objectives of this study were to determine: (i) the magnitude of diurnal and seasonal variation in CH₄ net flux, (ii) the relationship between the temporally varying flux rates and associated changes in controlling biotic and abiotic factors, and (iii) the contribution of CH₄ emission to the ecosystem growing season carbon budget. There was significant diurnal variation in CH₄ emission during the peak of the growing season that was strongly correlated with associated changes in solar radiation, latent heat flux, air temperature and ecosystem conductance to water vapor. During days 181–215, nighttime average CH₄ efflux was only 47% of the average midday values. The peak value for daily average CH₄ emission rate was approximately 80 nmol m^?2 s^?1 (4.6 mg CH₄ m^?2 h^?1), and seasonal variation in CH₄ flux was strongly correlated with changes in soil temperature. Integrated over the entire measurement period [days 144–269 (late May–late September)], the total CH₄ emission was 3.2 g CH₄ m^?2, which was quite low relative to other wetland ecosystems and to the simultaneous high rate of ecosystem net CO₂ sequestration that was measured (18.1 mol CO₂ m^?2 or 217 g C m^?2). We estimate that the negative radiative forcing (cooling) associated with net carbon storage over the life of the peatland (approximately 2200 years) was at least twice the value of positive radiative forcing (warming) caused by net CH₄ emission over the last 50 years.     

Controls on Soil Carbon Dioxide and Methane Fluxes in a Variety of Taiga Forest Stands in Interior Alaska

CO₂ and CH₄ fluxes were monitored over 4 years in a range of taiga forests along the Tanana River in interior Alaska. Floodplain alder and white spruce sites and upland birch/aspen and white spruce sites were examined. Each site had control, fertilized, and sawdust amended plots; flux measurements began during the second treatment year. CO₂ emissions decreased with successional age across the sites (alder, birch/aspen, and white spruce, in order of succession) regardless of landscape position. Although CO₂ fluxes showed an exponential relationship with soil temperature, the response of CO₂ production to moisture fit an asymptotic model. Of the manipulations, only N fertilization had an effect on CO₂ flux, decreasing flux in the floodplain sites but increasing it in the birch/aspen site. Landscape position was the best predictor of CH₄ flux. The two upland sites consumed CH₄ at similar rates (approximately 0.5 mg C m⁻² d⁻¹), whereas the floodplain sites had lower consumption rates (0–0.3 mg C m⁻² d⁻¹). N fertilization and sawdust both inhibited CH₄ consumption in the upland birch/aspen and floodplain spruce sites but not in the upland spruce site. The biological processes driving CO₂ fluxes were sensitive to temperature, moisture, and vegetation, whereas CH₄ fluxes were sensitive primarily to landscape position and biogeochemical disturbances. Hence, climate change effects on C-gas flux in taiga forest soils will depend on the relationship between soil temperature and moisture and the concomitant changes in soil nutrient pools and cycles. Received 10 March 1998; accepted 29 December 1999.     

Precipitation frequency alters peatland ecosystem structure and CO2 exchange: Contrasting effects on moss,sedge, and shrub communities

Climate projections forecast a redistribution of seasonal precipitation for much of the globe into fewer, larger events spaced between longer dry periods, with negligible changes in seasonal rainfall totals. This intensification of the rainfall regime is expected to alter near‐surface water availability, which will affect plant performance and carbon uptake. This could be especially important in peatland systems, where large stores of carbon are tightly coupled to water surpluses limiting decomposition. Here, we examined the role of precipitation frequency on vegetation growth and carbon dioxide (CO₂) balances for communities dominated by a Sphagnum moss, a sedge, and an ericaceous shrub in a cool temperate poor fen. Field plots and laboratory monoliths received one of three rainfall frequency treatments, ranging from one event every three days to one event every 14 days, while total rain delivered in a two‐week cycle and the entire season to each treatment remained the same. Separating incident rain into fewer but larger events increased vascular cover in all peatland communities: vascular plant cover increased 6× in the moss‐dominated plots, nearly doubled in the sedge plots, and tripled in the shrub plots in Low‐Frequency relative to High‐Frequency treatments. Gross ecosystem productivity was lowest in moss communities receiving low‐frequency rain, but higher in sedge and shrub communities under the same conditions. Net ecosystem exchange followed this pattern: fewer events with longer dry periods increased CO₂ flux to the atmosphere from the moss while vascular plant‐dominated communities became more of a sink for CO₂. Results of this study suggest that changes to rainfall frequency already occurring and predicted to continue will lead to increased vascular plant cover in peatlands and will impact their carbon‐sink function.     

Predominant role of water in regulating soil and microbial respiration and their responses to climate change in a semiarid grassland

Climate change can profoundly impact carbon (C) cycling of terrestrial ecosystems. A field experiment was conducted to examine responses of total soil and microbial respiration, and microbial biomass to experimental warming and increased precipitation in a semiarid temperate steppe in northern China since April 2005. We measured soil respiration twice a month over the growing seasons, soil microbial biomass C (MBC) and N (MBN), microbial respiration (MR) once a year in the middle growing season from 2005 to 2007. The results showed that interannual variations in soil respiration, MR, and microbial biomass were positively related to interannual fluctuations in precipitation. Laboratory incubation with a soil moisture gradient revealed a constraint of the temperature responses of MR by low soil moisture contents. Across the 3 years, experimental warming decreased soil moisture, and consequently caused significant reductions in total and microbial respiration, and microbial biomass, suggesting stronger negatively indirect effects through warming‐induced water stress than the positively direct effects of elevated temperature. Increased evapotranspiration under experimental warming could have reduced soil water availability below a stress threshold, thus leading to suppression of plant growth, root and microbial activities. Increased precipitation significantly stimulated total soil and microbial respiration and all other microbial parameters and the positive precipitation effects increased over time. Our results suggest that soil water availability is more important than temperature in regulating soil and microbial respiratory processes, microbial biomass and their responses to climate change in the semiarid temperate steppe. Experimental warming caused greater reductions in soil respiration than in gross ecosystem productivity (GEP). In contrast, increased precipitation stimulated GEP more than soil respiration. Our observations suggest that climate warming may cause net C losses, whereas increased precipitation may lead to net C gains in the semiarid temperate steppe. Our findings highlight that unless there is concurrent increase in precipitation, the temperate steppe in the arid and semiarid regions of northern China may act as a net C source under climate warming.     

Enhanced winter soil frost reduces methane emission during the subsequent growing season in a boreal peatland

Winter climate change may result in reduced snow cover and could, consequently, alter the soil frost regime and biogeochemical processes underlying the exchange of methane (CH₄) in boreal peatlands. In this study, we investigated the short‐term (1–3 years) vs. long‐term (11 years) effects of intensified winter soil frost (induced by experimental snow exclusion) on CH₄ exchange during the following growing season in a boreal peatland. In the first 3 years (2004–2006), lower CH₄ emissions in the treatment plots relative to the control coincided with delayed soil temperature increase in the treatment plots at the beginning of the growing season (May). After 11 treatment years (in 2014), CH₄ emissions were lower in the treatment plots relative to the control over the entire growing season, resulting in a reduction in total growing season CH₄ emission by 27%. From May to July 2014, reduced sedge leaf area coincided with lower CH₄ emissions in the treatment plots compared to the control. From July to August, lower dissolved organic carbon concentrations in the pore water of the treatment plots explained 72% of the differences in CH₄ emission between control and treatment. In addition, greater Sphagnum moss growth in the treatment plots resulted in a larger distance between the moss surface and the water table (i.e., increasing the oxic layer) which may have enhanced the CH₄ oxidation potential in the treatment plots relative to the control in 2014. The differences in vegetation might also explain the lower temperature sensitivity of CH₄ emission observed in the treatment plots relative to the control. Overall, this study suggests that greater soil frost, associated with future winter climate change, might substantially reduce the growing season CH₄ emission in boreal peatlands through altering vegetation dynamics and subsequently causing vegetation‐mediated effects on CH₄ exchange.     

Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes

The net primary productivity, carbon (C) stocks and turnover rates (i.e. C dynamics) of tropical forests are an important aspect of the global C cycle. These variables have been investigated in lowland tropical forests, but they have rarely been studied in tropical montane forests (TMFs). This study examines spatial patterns of above‐ and belowground C dynamics along a transect ranging from lowland Amazonia to the high Andes in SE Peru. Fine root biomass values increased from 1.50 Mg C ha^?1 at 194 m to 4.95 ± 0.62 Mg C ha^?1 at 3020 m, reaching a maximum of 6.83 ± 1.13 Mg C ha^?1 at the 2020 m elevation site. Aboveground biomass values decreased from 123.50 Mg C ha^?1 at 194 m to 47.03 Mg C ha^?1 at 3020 m. Mean annual belowground productivity was highest in the most fertile lowland plots (7.40 ± 1.00 Mg C ha^?1 yr^?1) and ranged between 3.43 ± 0.73 and 1.48 ± 0.40 Mg C ha^?1 yr^?1 in the premontane and montane plots. Mean annual aboveground productivity was estimated to vary between 9.50 ± 1.08 Mg C ha^?1 yr^?1 (210 m) and 2.59 ± 0.40 Mg C ha^?1 yr^?1 (2020 m), with consistently lower values observed in the cloud immersion zone of the montane forest. Fine root C residence time increased from 0.31 years in lowland Amazonia to 3.78 ± 0.81 years at 3020 m and stem C residence time remained constant along the elevational transect, with a mean of 54 ± 4 years. The ratio of fine root biomass to stem biomass increased significantly with increasing elevation, whereas the allocation of net primary productivity above‐ and belowground remained approximately constant at all elevations. Although net primary productivity declined in the TMF, the partitioning of productivity between the ecosystem subcomponents remained the same in lowland, premontane and montane forests.     

Spatially explicit patterns in a dryland's soil respiration and relationships with climate,whole plant photosynthesis and soil fertility

Arid and semiarid ecosystems play a significant role in regulating global carbon cycling, yet our understanding of the controls over the dominant pathways of dryland CO₂ exchange remains poor. Substantial amounts of dryland soil are not covered by vascular plants and this patchiness in cover has important implications for spatial patterns and controls of carbon cycling. Spatial variation in soil respiration has been attributed to variation in soil moisture, temperature, nutrients and rhizodeposition, while seasonal patterns have been attributed to changes in moisture, temperature and photosynthetic inputs belowground. To characterize how controls over respiration vary spatially and temporally in a dryland ecosystem and to concurrently explore multiple potential controls, we estimated whole plant net photosynthesis (A_net) and soil respiration at four distances from the plant base, as well as corresponding fine root biomass and soil carbon and nitrogen pools, four times during a growing season. To determine if the controls vary between different plant functional types for Colorado Plateau species, measurements were made on the C₄ shrub, Atriplex confertifolia, and C₃ grass, Achnatherum hymenoides. Soil respiration declined throughout the growing season and diminished with distance from the plant base, though variations in both were much smaller than expected. The strongest relationship was between soil respiration and soil moisture. Soil respiration was correlated with whole plant A_net, although the relationship varied between species and distance from plant base. In the especially dry year of this study we did not observe any consistent correlations between soil respiration and soil carbon or nitrogen pools. Our findings suggest that abiotic factors, especially soil moisture, strongly regulate the response of soil respiration to biotic factors and soil carbon and nitrogen pools in dryland communities and, at least in dry years, may override expected spatial and seasonal patterns.     

Soil CO2 effluxes,temporal and spatial variations,and root respiration in shrub willow biomass crops fields along a 19‐year chronosequence as affected by regrowth and removal treatments

In shrub willow biomass crop (SWBC) production systems, the soil CO₂ efflux (F_c) component in the carbon cycle remains poorly understood. This study assesses (i) differences of F_c rates among the 5‐, 12‐, 14‐, and 19‐year‐old SWBCs with two treatments: continuous production (regrowth) willow fields that were harvested and allowed to regrow, and willow fields that were harvested, killed, and then stools and roots were ground into the soil (removal); (ii) temporal and spatial variations of F_c rates; (iii) root respiration contributions to total F_c; and (iv) climatic variables affecting F_c. During the growing season (May to September), F_c rates showed no statistically significant differences across different ages (P = 0.664), and between treatments (P = 0.351); however, there was an interaction between age and treatment (P = 0.001). Similarly, during the dormant season (October to April), F_c rates revealed no statistically significant differences across different ages (P = 0.305) and treatment interaction with age (P = 0.097). F_c rates differed significantly (P < 0.001) among different times of the day and times of the year. F_c rates, between 00 and 1059 h, between 1100 and 1659 h, and between 1700 and 2400 h displayed consistency from May to November; however, F_c rates in these three time intervals showed significant differences (P < 0.0001). In December, F_c rates remained constant over 24 h. F_c rates demonstrated higher temporal and spatial variations among willow age classes than between regrowth and removal treatments. Temporal and spatial variations of F_c were higher during the dormant season than during the growing season. The proportion of root respiration to total F_c ranged from 18 to 33% across age classes. F_c rates showed strong association with soil and air temperatures, and relative humidity.     

Contrasting effects of Miocene and Anthropocene levels of atmospheric CO2 on silicon accumulation in a model grass

Grasses are hyper-accumulators of silicon (Si), which they acquire from the soil and deposit in tissues to resist environmental stresses. Given the high metabolic costs of herbivore defensive chemicals and structural constituents (e.g. cellulose), grasses may substitute Si for these components when carbon is limited. Indeed, high Si uptake grasses evolved in the Miocene when atmospheric CO₂ concentration was much lower than present levels. It is, however, unknown how pre-industrial CO₂ concentrations affect Si accumulation in grasses. Using Brachypodium distachyon, we hydroponically manipulated Si-supply (0.0, 0.5, 1, 1.5, 2 mM) and grew plants under Miocene (200 ppm) and Anthropocene levels of CO₂ comprising ambient (410 ppm) and elevated (640 ppm) CO₂ concentrations. We showed that regardless of Si treatments, the Miocene CO₂ levels increased foliar Si concentrations by 47% and 56% relative to plants grown under ambient and elevated CO₂, respectively. This is owing to higher accumulation overall, but also the reallocation of Si from the roots into the shoots. Our results suggest that grasses may accumulate high Si concentrations in foliage when carbon is less available (i.e. pre-industrial CO₂ levels) but this is likely to decline under future climate change scenarios, potentially leaving grasses more susceptible to environmental stresses.