Interaction of increasing atmospheric carbon dioxide and soil nitrogen on the carbon balance of tundra microcosms

Summary Natural cores of vegetation and soils of arctic tundra were collected in frozen condition in winter near Barrow, Alaska (71°20N). These cores were used as microcosms in a phytotron experiment to measure the interactions, if any, between increasing atmospheric CO₂ concentration and fertilization by ammonium nitrate on net ecosystem CO₂ exchange and net yield of tundra vegetation. Increased soil N significantly enhanced net ecosystem CO₂ uptake. The effect of increased CO₂ concentration had little or no effect on mean net ecosystem carbon balance of the tundra microcosms. Added N significantly increased leaf area and phytomass of vascular plants in the microcosms while increased atmospheric CO₂ had no effect on these parameters. We conclude that atmospheric CO₂ is not now limiting net ecosystem production in the tundra and that its direct effects will be slight even at double the present concentration. the most probable effects of carbon dioxide in the coastal tundra will be through its indirect effects on temperature, water table, peat decomposition, and the availability of soil nutrients.     

Effects of drainage and temperature on carbon balance of tussock tundra micrososms

We examined the importance of temperature (7°C or 15°C) and soil moisture regime (saturated or field capacity) on the carbon (C) balance of arctic tussock tundra microcosms (intact blocks of soil and vegetation) in growth chambers over an 81-day simulated growing season. We measured gaseous CO₂ exchanges, methane (CH₄) emissions, and dissolved C losses on intact blocks of tussock (Eriophorum vaginatum) and intertussock (moss-dominated). We hypothesized that under increased temperature and/or enhanced drainage, C losses from ecosystem respiration (CO₂ respired by plants and heterotrophs) would exceed gains from gross photosynthesis causing tussock tundra to become a net source of C to the atmosphere. The field capacity moisture regime caused a decrease in net CO₂ storage (NEP) in tussock tundra micrososms. This resulted from a stimulation of ecosystem respiration (probably mostly microbial) with enhanced drainage, rather than a decrease in gross photosynthesis. Elevated temperature alone had no effect on NEP because CO₂ losses from increased ecosystem respiration at elevated temperature were compensated by increased CO₂ uptake (gross photosynthesis). Although CO₂ losses from ecosystem respiration were primarily limited by drainage, CH₄ emissions, in contrast, were dependent on temperature. Furthermore, substantial dissolved C losses, especially organic C, and important microhabitat differences must be considered in estimating C balance for the tussock tundra system. As much as 20% of total C fixed in photosynthesis was lost as dissolved organic C. Tussocks stored 2x more C and emitted 5x more methane than intertussocks. In spite of the limitations of this microcosm experiment, this study has further elucidated the critical role of soil moisture regime and dissolved C losses in regulating net C balance of arctic tussock tundra.     

Carbon balance in tussock tundra under ambient and elevated atmospheric CO2

Summary Whole ecosystem CO₂ flux under ambient (340 l/l) and elevated (680 l/l) CO₂ was measured in situ in Eriophorum tussock tundra on the North Slope of Alaska. Elevated CO₂ resulted in greater carbon acquisition than control treatments and there was a net loss of CO₂ under ambient conditions at this upland tundra site. These measurements indicate a current loss of carbon from upland tundra, possibly the result of recent climatic changes. Elevated CO₂ for the duration of one growing season appeared to delay the onset of dormancy and resulted in approximately 10 additional days of positive ecosystem flux. Homeostatic adjustment of ecosystem CO₂ flux (sum of species' response) was apparent by the third week of exposure to elevated CO₂. Ecosystem dark respiration rates were not significantly higher at elevated CO₂ levels. Rapid homeostatic adjustment to elevated CO₂ may limit carbon uptake in upland tundra. Abiotic factors were evaluated as predictors of ecosystem CO₂ flux. For chambers exposed to ambient and elevated CO₂ levels for the duration of the growing season, seasonality (Julian day) was the best predictor of ecosystem CO₂ flux at both ambient and elevated CO₂ levels. Light (PAR), soil temperature, and air temperature were also predictive of seasonal ecosystem flux, but only at elevated CO₂ levels. At any combination of physical conditions, flux of the elevated CO₂ treatment was greater than that at ambient. In short-term manipulations of CO₂, tundra exposed to elevated CO₂ had threefold greater carbon gain, and had one half the ecosystem level, light compensation point when compared to ambient CO₂ treatments. Elevated CO₂-acclimated tundra had twofold greater carbon gain compared to ambient treatments, but there was no difference in ecosystem level, light compensation point between elevated and ambient CO₂ treatments. The predicted future increases in cloudiness could substantially decrease the effect of elevated atmospheric CO₂ on net ecosystem carbon budget. These analyses suggest little if any long-term stimulation of ecosystem carbon acquisition by increases in atmospheric CO₂.     

Methanogenesis at low temperatures by microflora of tundra wetland soil

Active methanogenesis from organic matter contained in soil samples from tundra wetland occurred even at 6 °C. Methane was the only end product in balanced microbial community with H₂/CO₂ as a substrate, besides acetate was produced as an intermediate at temperatures below 10°C. The activity of different microbial groups of methanogenic community in the temperature range of 6–28 °C was investigated using 5% of tundra soil as inoculum. Anaerobic microflora of tundra wetland fermented different organic compounds with formation of hydrogen, volatile fatty acids (VFA) and alcohols. Methane was produced at the second step. Homoacetogenic and methanogenic bacteria competed for such substrates as hydrogen, formate, carbon monoxide and methanol. Acetogens out competed methanogens in an excess of substrate and low density of microbial population. Kinetic analysis of the results confirmed the prevalence of hydrogen acetogenesis on methanogenesis. Pure culture of acetogenic bacteria was isolated at 6 °C. Dilution of tundra soil and supply with the excess of substrate disbalanced the methanoigenic microbial community. It resulted in accumulation of acetate and other VFA. In balanced microbial community obviously autotrophic methanogens keep hydrogen concentration below a threshold for syntrophic degradation of VFA. Accumulation of acetate- and H₂/CO₂-utilising methanogens should be very important in methanogenic microbial community operating at low temperatures.     

Biogeochemical pools and fluxes of carbon and nitrogen in a maritime tundra near penguin colonies along the Antarctic Peninsula

There is limited information regarding biogeochemical pools and fluxes in maritime tundra ecosystems along the Antarctic Peninsula. To collect baseline information on biogeochemical processes in a tundra ecosystem dominated by two vascular plant species (Colobanthus quitensis and Deschampsia antarctica) at Biscoe Point off the coast of Anvers Island, we measured pools and fluxes of C and N in transplanted tundra microcosm cores, complemented with sampling of precipitation and surface runoff. Snow and snowmelt from the tundra collection site and soil leachates from the cores were enriched with N and dissolved organic carbon compared to precipitation and snowmelt samples collected at Palmer Station, indicating high loading of N and organic matter from the penguin colonies adjacent to the tundra site. Relatively high values of δ¹⁵N in the live and dead biomass of D. antarctica and C. quitensis (5.6–25.1‰) indicated an enrichment of N in this tundra ecosystem, possibly through N inputs from adjacent penguin colonies. Stepwise multiple linear regressions found that ecosystem respiration and gross primary production were best predicted by live biomass of D. antarctica, suggesting a disproportionately high contribution of D. antarctica to CO₂ fluxes. The cores with higher δ¹⁵N and lower δ¹³C in the soil organic horizon exhibited higher CO₂ fluxes. The results suggest that abundant N inputs from penguin colonies and the competitive balance between plant species might play a critical role in the response of tundra ecosystems along the Antarctic Peninsula to projected climate change.     

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.     

Simulation of tree species composition and organic matter accumulation in Finnish boreal forests under changing climatic conditions

A model simulating the regeneration, growth and death of trees and the consequent carbon and nitrogen dynamics of the forest ecosystem was applied to determine the effect of expected temperature rise on tree species composition and the accumulation of organic matter in the boreal forest ecosystem in Finland (between latitudes 60°–70° N). In the southern and middle boreal zones a temperature rise of 2–3° C (temperature for 2 x CO₂) over a period of one hundred years increased the competitive capacity of Scots pine (Pinus sylvestris) and birch species (Betula pendula and B. pubescens), and slowed down the invasion by Norway spruce (Picea abies). In the northern boreal zone a corresponding rise in temperature promoted the invasion of sites by Norway spruce. The accumulation of organic matter was promoted only slightly compared to that taking place in the current climatic conditions.A further doubling of temperature (temperature for 4 x CO₂) over an additional period of two hundred years led to the replacement of coniferous stands with deciduous onesin the southern and middle boreal zones. In the northern boreal zone an admixture of coniferous and deciduous species replaced pure coniferous stands with the latter taking over sites formerly classified as tundra woodland. In the southern and middle boreal zones the replacement of coniferous species induced a substantial decrease in the amount of organic matter; this returned to its former level following the establishment of deciduous species. In the northern boreal zone there was no major change in the amount of organic matter such as occurred in the case of the tundra woodland where the amount of organic matter accumulated was nearly as high as in the northern boreal zone.     

Long-Term Release of Carbon Dioxide from Arctic Tundra Ecosystems in Alaska

Releases of the greenhouse gases carbon dioxide (CO₂) and methane (CH₄) from thawing permafrost are expected to be among the largest feedbacks to climate from arctic ecosystems. However, the current net carbon (C) balance of terrestrial arctic ecosystems is unknown. Recent studies suggest that these ecosystems are sources, sinks, or approximately in balance at present. This uncertainty arises because there are few long-term continuous measurements of arctic tundra CO₂ fluxes over the full annual cycle. Here, we describe a pattern of CO₂ loss based on the longest continuous record of direct measurements of CO₂ fluxes in the Alaskan Arctic, from two representative tundra ecosystems, wet sedge and heath tundra. We also report on a shorter time series of continuous measurements from a third ecosystem, tussock tundra. The amount of CO₂ loss from both heath and wet sedge ecosystems was related to the timing of freeze-up of the soil active layer in the fall. Wet sedge tundra lost the most CO₂ during the anomalously warm autumn periods of September–December 2013–2015, with CH₄ emissions contributing little to the overall C budget. Losses of C translated to approximately 4.1 and 1.4% of the total soil C stocks in active layer of the wet sedge and heath tundra, respectively, from 2008 to 2015. Increases in air temperature and soil temperatures at all depths may trigger a new trajectory of CO₂ release, which will be a significant feedback to further warming if it is representative of larger areas of the Arctic.     

Temperature response of CO<Subscript>2</Subscript> exchange and dissolved organic carbon release in a maritime Antarctic tundra ecosystem

We examined the temperature response of CO₂ exchange and soil biogeochemical processes in an Antarctic tundra ecosystem using laboratory incubations of intact tundra cores. The cores were collected from tundra near Anvers Island along the west coast of the Antarctic Peninsula that was dominated by the vascular plants Colobanthus quitensis and Deschampsia antarctica. After the initial 8-week incubation at moderate growth temperatures (12/7°C, day/night), the tundra cores were incubated for another 8 weeks at either a higher (17/12°C) or lower (7/4°C) temperature regime. Temperature responses of CO₂ exchange were measured at five temperatures (4, 7, 12, 17, and 27°C) following each incubation and soil leachates were collected biweekly over the second incubation. Daytime net ecosystem CO₂ exchange (NEE) per unit core surface area was higher across the five measurement temperatures after the warmer incubation (17/12°C > 7/4°C). Responses of ecosystem respiration (ER) were similar at each measurement temperature irrespective of incubation temperature regimes. ER, expressed on a leaf-area basis, however, was significantly lower following the warmer incubation, suggesting a downregulation of ER. Warmer incubation resulted in a greater specific leaf area and N concentration, and a lower δ¹³C in live aboveground C. quitensis, but a higher δ¹³C in D. antarctica, implying species-specific responses to warming. Concentrations of dissolved organic C and N and inorganic N in soil leachates showed that short-term temperature changes had no noticeable effect on soil biogeochemical processes. The results suggest that downregulation of ER, together with plant species differences in leaf-area production and N use, can play a crucial role in constraining the C-cycle response of Antarctic tundra ecosystems to warming.     

Spring photosynthetic onset and net CO2 uptake in Alaska triggered by landscape thawing

The springtime transition to regional‐scale onset of photosynthesis and net ecosystem carbon uptake in boreal and tundra ecosystems are linked to the soil freeze–thaw state. We present evidence from diagnostic and inversion models constrained by satellite fluorescence and airborne CO₂ from 2012 to 2014 indicating the timing and magnitude of spring carbon uptake in Alaska correlates with landscape thaw and ecoregion. Landscape thaw in boreal forests typically occurs in late April (DOY 111 ± 7) with a 29 ± 6 day lag until photosynthetic onset. North Slope tundra thaws 3 weeks later (DOY 133 ± 5) but experiences only a 20 ± 5 day lag until photosynthetic onset. These time lag differences reflect efficient cold season adaptation in tundra shrub and the longer dehardening period for boreal evergreens. Despite the short transition from thaw to photosynthetic onset in tundra, synchrony of tundra respiration with snow melt and landscape thaw delays the transition from net carbon loss (at photosynthetic onset) to net uptake by 13 ± 7 days, thus reducing the tundra net carbon uptake period. Two global CO₂ inversions using a CASA‐GFED model prior estimate earlier northern high latitude net carbon uptake compared to our regional inversion, which we attribute to (i) early photosynthetic‐onset model prior bias, (ii) inverse method (scaling factor + optimization window), and (iii) sparsity of available Alaskan CO₂ observations. Another global inversion with zero prior estimates the same timing for net carbon uptake as the regional model but smaller seasonal amplitude. The analysis of Alaskan eddy covariance observations confirms regional scale findings for tundra, but indicates that photosynthesis and net carbon uptake occur up to 1 month earlier in evergreens than captured by models or CO₂ inversions, with better correlation to above‐freezing air temperature than date of primary thaw. Further collection and analysis of boreal evergreen species over multiple years and at additional subarctic flux towers are critically needed.     

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.     

Soil respiration of Alaskan tundra at elevated atmospheric carbon dioxide concentrations

Summary CO₂ efflux from tussock tundra in Alaska that had been exposed to elevated CO₂ for 2.5 growing seasons was measured to assess the effect of long- and short-term CO₂ enrichment on soil respiration. Long-term treatments were: 348, 514, and 683 μll⁻¹ CO₂ and 680 μll⁻¹ CO₂+4°C above ambient. Measurements were made at 5 CO₂ concentrations between 87 and 680 μll⁻¹ CO₂. Neither long- or short-term CO₂ enrichment significantly affected soil CO₂ efflux. Tundra developed at elevated temperature and 680 μll⁻¹ CO₂ had slightly higher, but not statistically different, mean respiration rates compared to untreated tundra and to tundra under CO₂ control alone.     

A scaling approach for quantifying the net CO2 flux of the Kuparuk River Basin,Alaska

Net CO₂ flux measurements conducted during the summer and winter of 1994–96 were scaled in space and time to provide estimates of net CO₂ exchange during the 1995–96 (9 May 1995–8 May 1996) annual cycle for the Kuparuk River Basin, a 9200 km² watershed located in NE Alaska. Net CO₂ flux was measured using dynamic chambers and eddy covariance in moist‐acidic, nonacidic, wet‐sedge, and shrub tundra, which comprise 95% of the terrestrial landscape of the Kuparuk Basin. CO₂ flux data were used as input to multivariate models that calculated instantaneous and daily rates of gross primary production (GPP) and whole‐ecosystem respiration (R) as a function of meteorology and ecosystem development. Net CO₂ flux was scaled up to the Kuparuk Basin using a geographical information system (GIS) consisting of a vegetation map, digital terrain map, dynamic temperature and radiation fields, and the models of GPP and R. Basin‐wide estimates of net CO₂ exchange for the summer growing season (9 May?5 September 1995) indicate that nonacidic tundra was a net sink of ?31.7 ± 21.3 GgC (1 Gg = 10⁹ g), while shrub tundra lost 32.5 ± 6.3 GgC to the atmosphere (negative values denote net ecosystem CO₂ uptake). Acidic and wet sedge tundra were in balance, and when integrated for the entire Kuparuk River Basin (including aquatic surfaces), whole basin summer net CO₂ exchange was estimated to be in balance (?0.9 ± 50.3 GgC). Autumn to winter (6 September 1995–8 May 1996) estimates of net CO₂ flux indicate that acidic, nonacidic, and shrub tundra landforms were all large sources of CO₂ to the atmosphere (75.5 ± 8.3, 96.4 ± 11.4, and 43.3 ± 4.7 GgC for acidic, nonacidic, and shrub tundra, respectively). CO₂ loss from wet sedge surfaces was not substantially different from zero, but the large losses from the other terrestrial landforms resulted in a whole basin net CO₂ loss of 217.2 ± 24.1 GgC during the 1995–96 cold season. When integrated for the 1995–96 annual cycle, acidic (66.4 + 25.25 GgC), nonacidic (64.7 ± 29.2 GgC), and shrub tundra (75.8 ± 8.4 GgC) were substantial net sources of CO₂ to the atmosphere, while wet sedge tundra was in balance (0.4 + 0.8 GgC). The Kuparuk River Basin as a whole was estimated to be a net CO₂ source of 218.1 ± 60.6 GgC over the 1995–96 annual cycle. Compared to direct measurements of regional net CO₂ flux obtained from aircraft‐based eddy covariance, the scaling procedure provided realistic estimates of CO₂ exchange during the summer growing season. Although winter estimates could not be assessed directly using aircraft measurements of net CO₂ exchange, the estimates reported here are comparable to measured values reported in the literature. Thus, we have high confidence in the summer estimates of net CO₂ exchange and reasonable confidence in the winter net CO₂ flux estimates for terrestrial landforms of the Kuparuk river basin. Although there is larger uncertainty in the aquatic estimates, the small surface area of aquatic surfaces in the Kuparuk river basin (≈ 5%) presumably reduces the potential for this uncertainty to result in large errors in basin‐wide CO₂ flux estimates.     

Snow distribution, soil temperature and late winter CO2 efflux from soils near the Arctic treeline in northwest Alaska

The Arctic treeline is advancing in many areas and changes in carbon (C) cycling are anticipated. Differences in CO₂ exchange between adjacent forest and tundra are not well known and contrasting conclusions have been drawn about the effects of forest advance on ecosystem C stocks. Measurements of CO₂ exchange in tundra and adjacent forest showed the forest was a greater C sink during the growing season in northern Canada. There is, however, reason to expect that forests lose more C than tundra during the wintertime, as forests may accumulate and retain more snow. Deeper snow insulates the soil and warmer soils should lead to greater rates of belowground respiration and CO₂ efflux. In this study, I tested the hypotheses that forests maintain a deeper snowpack, have warmer soils and lose more C during winter than adjacent tundra near the Arctic treeline in northwest Alaska. Measurements of snow depth, soil temperature and CO₂ efflux were made at five forest and two treeline sites in late winter of three consecutive years. Snow depth and soil temperature were greater in forest than treeline sites, particularly in years with higher snowfall. There was a close exponential correlation between soil temperature and CO₂ efflux across sites and years. The temperature-efflux model was driven using hourly soil temperatures from all the sites to provide a first approximation of the difference in winter C loss between treeline and forest sites. Results showed that greater wintertime C loss from forests could offset greater summertime C gain.     

The subdominant status of Echinocereus viridiflorus and Mammillaria vivipara in the shortgrass prairie: The role of temperature and water effects on gas exchange

Summary The subdominant CAM species, Echinocereus viridiflorus and Mammillaria vivipara, collected from the shortgrass prairie in northeastern Colorado were pretreated and analyzed for gas exchange under cool temperatures (20/15°C) and warm temperatures (35/15°C). Well watered plants of both species under a 35/15°C thermoperiod fixed atmospheric CO₂ during the night and early moring. Echinocereus viridiflorus grown and analyzed at 20/15°C fixed CO₂ during the night, early morning and late afternoon but total carbon gain over a 24 h period is less than when grown and analyzed under the 35/15°C thermoperiod. Mammillaria vivipara grown and analyzed at 20/15°C assimilates CO₂ at low rates during all parts of a 24 h period with the greatest CO₂ fixation rates occuring from midday to late afternoon. The total carbon gain under the 20/15°C thermoperiod is less than that for this species under the 35/15°C thermoperiod. Decreasing the night temperature of plants grown under the warm conditions to 10°C or 5°C results in a depression of the night CO₂ fixation in both species. E. viridiflorus from the cool growth conditions showed an enhancement of the CO₂ uptake during the night, early morning and late afternoon when subjected to the cooler night temperatures (10°C and 5°C). The CO₂ uptake of M. vivipara grown at 20/15°C shows an enhancement during the night and early morning while the CO₂ fixation during midday and late afternoon is slightly depressed under cool night temperatures (10° and 5°C). Under the 35/15°C thermoperiod both species exhibit depressed rates of CO₂ fixation during the night and early morning when water stressed. Plants of both species grown under the 20/15°C thermoperiod exhibit no net CO₂ fixation following five weeks of water deprivation. Upon rewatering, E. viridiflorus begins to recover its capacity for CO₂ fixation within 24 h under both the warm and cool temperature regimes. However, M. vivipara did not show recovery within 48 h following rewatering under the warm or cool temperature regime. Contrasting the patterns of gas exchange of the subdominant species, E. viridiflorus and M. vivipara, with a dominant CAM species of the shortgrass prairie, Opuntia polyacantha reveals significant differences that may well dictate the role of these species in this ecosystem. E. viridiflorus and M. vivipara have a lower capacity of carbon gain and recovery from water stress than O. polyacantha mainly due to their lack of late afternoon CO₂ uptake. This study suggests that carbon gain plays an important role in limiting E. viridiflorus and M. vivipara in the shortgrass prairie ecosystem.     

Tundra ecosystems observed to be CO2 sources due to differential amplification of the carbon cycle

Are tundra ecosystems currently a carbon source or sink? What is the future trajectory of tundra carbon fluxes in response to climate change? These questions are of global importance because of the vast quantities of organic carbon stored in permafrost soils. In this meta‐analysis, we compile 40 years of CO₂ flux observations from 54 studies spanning 32 sites across northern high latitudes. Using time‐series analysis, we investigated if seasonal or annual CO₂ fluxes have changed over time, and whether spatial differences in mean annual temperature could help explain temporal changes in CO₂ flux. Growing season net CO₂ uptake has definitely increased since the 1990s; the data also suggest (albeit less definitively) an increase in winter CO₂ emissions, especially in the last decade. In spite of the uncertainty in the winter trend, we estimate that tundra sites were annual CO₂ sources from the mid‐1980s until the 2000s, and data from the last 7 years show that tundra continue to emit CO₂ annually. CO₂ emissions exceed CO₂ uptake across the range of temperatures that occur in the tundra biome. Taken together, these data suggest that despite increases in growing season uptake, tundra ecosystems are currently CO₂ sources on an annual basis.     

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.     

Environmental and Vegetation Drivers of Seasonal CO2 Fluxes in a Sub-arctic Forest–Mire Ecotone

Unravelling the role of structural and environmental drivers of gross primary productivity (GPP) and ecosystem respiration (R _eco) in highly heterogeneous tundra is a major challenge for the upscaling of chamber-based CO₂ fluxes in Arctic landscapes. In a mountain birch woodland-mire ecotone, we investigated the role of LAI (and NDVI), environmental factors (microclimate, soil moisture), and microsite type across tundra shrub plots (wet hummocks, dry hummocks, dry hollows) and lichen hummocks, in controlling net ecosystem CO₂ exchange (NEE). During a growing season, we measured NEE fluxes continuously, with closed dynamic chambers, and performed multiple fits (one for each 3-day period) of a simple light and temperature response model to hourly NEE data. Tundra shrub plots were largely CO₂ sinks, as opposed to lichen plots, although fluxes were highly variable within microsite type. For tundra shrub plots, microsite type did not influence photosynthetic parameters but it affected basal (that is, temperature-normalized) ecosystem respiration (R ₀). PAR-normalized photosynthesis (P ₆₀₀) increased with air temperature and declined with increasing vapor pressure deficit. R ₀ declined with soil moisture and showed an apparent increase with temperature, which may underlie a tight link between GPP and R _eco. NDVI was a good proxy for LAI, maximum P ₆₀₀ and maximum R ₀ of shrub plots. Cumulative CO₂ fluxes were strongly correlated with LAI (NDVI) but we observed a comparatively low GPP/LAI in dry hummocks. Our results broadly agree with the reported functional convergence across tundra vegetation, but here we show that the role of decreased productivity in transition zones and the influence of temperature and water balance on seasonal CO₂ fluxes in sub-Arctic forest–mire ecotones cannot be overlooked.     

Carbon balance in the tundra, boreal forest and humid tropical forest during climate change: scaling up from leaf physiology and soil carbon dynamics

Carbon exchange by the terrestrial biosphere is thought to have changed since pre-industrial times in response to increasing concentrations of atmospheric CO₂ and variations (anomalies) in inter-annual air temperatures. However, the magnitude of this response, particularly that of various ecosystem types (biomes), is uncertain. Terrestrial carbon models can be used to estimate the direction and size of the terrestrial responses expected, providing that these models have a reasonable theoretical base. We formulated a general model of ecosystem carbon fluxes by linking a process-based canopy photosynthesis model to the Rothamsted soil carbon model for biomes that are not significantly affected by water limitation. The difference between net primary production (NPP) and heterotrophic soil respiration (R_h) represents net ecosystem production (NEP). The model includes (i) multiple compartments for carbon storage in vegetation and soil organic matter, (ii) the effects of seasonal changes in environmental parameters on annual NEP, and (iii) the effects of inter-annual temperature variations on annual NEP. Past, present and projected changes in atmospheric CO₂ concentration and surface air temperature (at different latitudes) were analysed for their effects on annual NEP in tundra, boreal forest and humid tropical forest biomes. In all three biomes, annual NEP was predicted to increase with CO₂ concentration but to decrease with warming. As CO₂ concentrations and temperatures rise, the positive carbon gains through increased NPP are often outweighed by losses through increased R_h, particularly at high latitudes where global warming has been (and is expected to be) most severe. We calculated that, several times during the past 140 years, both the tundra and boreal forest biomes have switched between being carbon sources (annual NEP negative) and being carbon sinks (annual NEP positive). Most recently, significant warming at high latitudes during 1988 and 1990 caused the tundra and boreal forests to be net carbon sources. Humid tropical forests generally have been a carbon sink since 1960. These modelled responses of the various biomes are in agreement with other estimates from either field measurements or geochemical models. Under projected CO₂ and temperature increases, the tundra and boreal forests will emit increasingly more carbon to the atmosphere while the humid tropical forest will continue to store carbon. Our analyses also indicate that the relative increase in the seasonal amplitude of the accumulated NEP within a year is about 0–14% year^?1 for boreal forests and 0–23% year^?1 in the tundra between 1960 and 1990.     

Temperature dependence of CO2 assimilation and stomatal aperture in leaf sections of Zea mays

Summary CO₂ exchange and air flow through the stomata were measured in leaf sections of Zea mays at temperatures between 7 and 52° and under optimal water supply. The results were summarized in polynomials fitted to the data.In leaf samples brought from 16° and darkness into different experimental temperatures and light, CO₂ assimilation has a maximum near 30°. Above 37° (in other experiments above 41°), net CO₂ uptake stops abruptly and is replaced by CO₂ evolution in light. If a 1-hr treatment with 25° and light is inserted between darkness and the experimental temperatures, the threshold above which the assimilatory system collapses shifts 3 degrees upwards, to 40° (or 44°); the decline of CO₂ assimilation with high temperatures is less steep than without pretreatment; and the upper compensation point moves upscale by as much as 5 degrees.Stomatal conductance for CO₂ does not, in general, follow an optimum curve with temperature. Between 15 and 35° it is approximately proportional to net CO₂ assimilation, indicating control by CO₂; but above 35°, stomatal aperture increases further with temperature (and so does stomatal variability): the stomata escape the control by CO₂ and above 40° may be wide open even if CO₂ is being evolved. Stomatal conductance for CO₂ below 15° may also be larger than would be proportional to CO₂ assimilation.Net CO₂ assimilation and stomatal conductance at 25° were reduced if the leaf samples were pretreated with temperatures below approximately 20° and above 30°. Stomata were more sensitive to past temperatures than was CO₂ assimilation.