Variations of ecosystem gas exchange in the rain forest mesocosm at Biosphere 2 in response to elevated CO2

The effects of elevated CO₂ on tropical ecosystems were studied in the artificial rain forest mesocosm at Biosphere 2, a large-scale and ecologically diverse experimental facility located in Oracle, Arizona. The ecosystem responses were assessed by comparing the whole-system net gas exchange (NEE) upon changing CO₂ levels from 900 to 450 ppmV. The day-NEE was significantly higher in the elevated CO₂ treatment. In both experiments, the NEE rates were similar to values observed in natural analogue systems. Variations in night-NEE, reflecting both soil CO₂ efflux and plants respiration, covaried with temperature but showed no clear correlation with atmospheric CO₂ levels. After correcting for changes in CO₂ efflux we show that the rain forest net photosynthesis increased in response to increasing atmospheric CO₂. The photosynthetic enhancement was expressed in higher quantum yields, maximum assimilation rates and radiation use efficiency. The results suggest that photosynthesis in large tropical trees is CO₂ sensitive, at least following short exposures of days to weeks. Taken at face value, the data suggest that as a result of anthropogenic emissions of CO₂, tropical rain forests may shift out of steady state, and become a carbon sink at least for short periods. However, a better understanding of the unique conditions and phenomena in Biosphere 2 is necessary before these results are broadly useful.     

The contribution of bryophytes to the carbon exchange for a temperate rainforest

Bryophytes blanket the floor of temperate rainforests in New Zealand and may influence a number of important ecosystem processes, including carbon cycling. Their contribution to forest floor carbon exchange was determined in a mature, undisturbed podocarp‐broadleaved forest in New Zealand, dominated by 100–400‐year‐old rimu (Dacrydium cupressimum) trees. Eight species of mosses and 13 species of liverworts contributed to the 62% cover of the diverse forest floor community. The bryophyte community developed a relatively thin (depth <30 mm), but dense, canopy that experienced elevated CO₂ partial pressures (median 46.6 Pa immediately below the bryophyte canopy) relative to the surrounding air (median 37.6 Pa at 100 mm above the canopy). Light‐saturated rates of net CO₂ exchange from 14 microcosms collected from the forest floor were highly variable; the maximum rate of net uptake (bryophyte photosynthesis – whole‐plant respiration) per unit ground area at saturating irradiance was 1.9 μmol m^?2 s^?1 and in one microcosm, the net rate of CO₂ exchange was negative (respiration). CO₂ exchange for all microcosms was strongly dependent on water content. The average water content in the microcosms ranged from 1375% when fully saturated to 250% when air‐dried. Reduction in water content across this range resulted in an average decrease of 85% in net CO₂ uptake per unit ground area. The results from the microcosms were used in a model to estimate annual carbon exchange for the forest floor. This model incorporated hourly variability in average irradiance reaching the forest floor, water content of the bryophyte layer, and air and soil temperature. The annual net carbon uptake by forest floor bryophytes was 103 g m^?2, compared to annual carbon efflux from the forest floor (bryophyte and soil respiration) of ?1010 g m^?2. To put this in perspective of the magnitude of the components of CO₂ exchange for the forest floor, the bryophyte layer reclaimed an amount of CO₂ equivalent to only about 10% of forest floor respiration (bryophyte plus soil) or ～11% of soil respiration. The contribution of forest floor bryophytes to productivity in this temperate rainforest was much smaller than in boreal forests, possibly because of differences in species composition and environmental limitations to photosynthesis. Because of their close dependence on water table depth, the contribution of the bryophyte community to ecosystem CO₂ exchange may be highly responsive to rapid changes in climate.     

Modelling night‐time ecosystem respiration by a constrained source optimization method

One of the main challenges to quantifying ecosystem carbon budgets is properly quantifying the magnitude of night‐time ecosystem respiration. Inverse Lagrangian dispersion analysis provides a promising approach to addressing such a problem when measured mean CO₂ concentration profiles and nocturnal velocity statistics are available. An inverse method, termed ‘Constrained Source Optimization’ or CSO, which couples a localized near‐field theory (LNF) of turbulent dispersion to respiratory sources, is developed to estimate seasonal and annual components of ecosystem respiration. A key advantage to the proposed method is that the effects of variable leaf area density on flow statistics are explicitly resolved via higher‐order closure principles. In CSO, the source distribution was computed after optimizing key physiological parameters to recover the measured mean concentration profile in a least‐square fashion. The proposed method was field‐tested using 1 year of 30‐min mean CO₂ concentration and CO₂ flux measurements collected within a 17‐year‐old (in 1999) even‐aged loblolly pine (Pinus taeda L.) stand in central North Carolina. Eddy‐covariance flux measurements conditioned on large friction velocity, leaf‐level porometry and forest‐floor respiration chamber measurements were used to assess the performance of the CSO model. The CSO approach produced reasonable estimates of ecosystem respiration, which permits estimation of ecosystem gross primary production when combined with daytime net ecosystem exchange (NEE) measurements. We employed the CSO approach in modelling annual respiration of above‐ground plant components (c. 214 g C m^?2 year^?1) and forest floor (c. 989 g C m^?2 year^?1) for estimating gross primary production (c. 1800 g C m^?2 year^?1) with a NEE of c. 605 g C m^?2 year^?1 for this pine forest ecosystem. We conclude that the CSO approach can utilise routine CO₂ concentration profile measurements to corroborate forest carbon balance estimates from eddy‐covariance NEE and chamber‐based component flux measurements.     

Carbon balance of different aged Scots pine forests in Southern Finland

We estimated annual net ecosystem exchange (NEE) of a chronosequence of four Scots pine stands in southern Finland during years 2000–2002 using eddy covariance (EC). Net ecosystem productivity (NEP) was estimated using growth measurements and modelled mass losses of woody debris. The stands were 4, 12, 40 and 75 years old. The 4‐year‐old clearcut was a source of carbon throughout the year combining a low gross primary productivity (GPP) with a total ecosystem respiration (TER) similar to the forest stands. The annual NEE of the clearcut, measured by EC, was 386 g C m^?2. Tree growth was negligible and the estimated NEP was ?262 g C m^?2 a^?1. The annual GPPs at the other sites were close to each other (928?1072 g C m^?2 a^?1), but TER differed markedly, being greatest at the 12‐year‐old site (905 g C m^?2 a^?1) and smallest in the 75‐year‐old stand (616 g C m^?2 a^?1). Measurements of soil CO₂ efflux showed that different rates of soil respiration largely explained the differences in TER. The NEE and NEP of the 12‐year‐old stand were close to zero. The forested stands were sinks of carbon. They had similar annual patterns of carbon exchange and half‐hourly eddy fluxes were highly correlated, indicating similar responses to the environment. The NEE in the 40‐year‐old stand varied between ?179 and –192 g C m^?2 a^?1, while NEP was between 214 and 242 g C m^?2 a^?1. The annual NEE of the 75‐year‐old stand was 323 g C m^?2 and NEP was 252 g C m^?2. This indicates that there was no reduction in carbon sink strength with stand age.     

Annual cycles of water vapour and carbon dioxide fluxes in and above a boreal aspen forest

Water vapour and CO₂ fluxes were measured using the eddy correlation method above and below the overstorey of a 21-m tall aspen stand in the boreal forest of central Saskatchewan as part of the Boreal Ecosystem-Atmosphere Study (BOREAS). Measurements were made at the 39.5-m and 4-m heights using 3-dimensional sonic anemometers (Kaijo-Denki and Solent, respectively) and closed-path gas analysers (LI-COR 6262) with 6-m and 4.7-m long heated sampling tubing, respectively. Continuous measurements were made from early October to mid-November 1993 and from early February to late-September 1994. Soil CO₂ flux (respiration) was measured using a LI-COR 6000-09 soil chamber and soil evaporation was measured using Iysimetry. The leaf area index of the aspen and hazelnut understorey reached 1.8 and 3.3, respectively. The maximum daily evapotranspiration (E) rate was 5–6 mm d^?1. Following leaf-out the hazelnut and soil accounted for 22% of the forest E. The estimated total E was 403 mm for 1994. About 88% of the precipitation in 1994 was lost as evapotranspiration. During the growing season, the magnitude of half-hourly eddy fluxes of CO₂ from the atmosphere into the forest reached 1.2 mg CO₂ m^?2 s^?1 (33 μmol C m^?2 s^?1) during the daytime. Downward eddy fluxes at the 4-m height were observed when the hazelnut was growing rapidly in June and July. Under well-ventilated night-time conditions, the eddy fluxes of CO₂ above the aspen and hazelnut, corrected for canopy storage, increased exponentially with soil temperature at the 2-cm depth. Estimates of daytime respiration rates using these relationships agreed well with soil chamber measurements. During the 1994 growing season, the cumulative net ecosystem exchange (NEE) was -3.5 t C ha^?1 y^?1 (a net gain by the system). For 1994, cumulative NEE, ecosystem respiration (R) and gross ecosystem photosynthesis (GEP = R - NEE) were estimated to be -1.3, 8.9 and 10.2 t C ha^?1 y^?1 respectively. Gross photosynthesis of the hazelnut was 32% of GEP.     

Carbon sequestration in boreal jack pine stands following harvesting

A large area of boreal jack pine (Pinus banksiana Lamb.) forest in Canada is recovering from clear‐cut harvesting, and the carbon (C) balance of these regenerating forests remains uncertain. Net ecosystem CO₂ exchange was measured using the eddy‐covariance technique at four jack pine sites representing different stages of stand development: three postharvest sites (HJP02, HJP94, and HJP75) and one preharvest site (OJP). The four sites, located in the southern Canadian boreal forest, Saskatchewan, Canada, are typical of low productivity jack pine stands and were 2, 10, 29, and 90 years old in 2004, respectively. Mean annual net ecosystem production (NEP) for 2004 and 2005 was ?137±11, 19±16, 73±28, and 22±30 g C m^?2 yr^?1 at HJP02, HJP94, HJP75 and OJP, respectively, showing the postharvest jack pine stands to be moderate C sources immediately after harvesting, weak sinks at 10 years, moderate C sinks at 30 years, then weak C sinks at 90 years. Mean annual gross ecosystem photosynthesis (GEP) for the 2 years was 96±10, 347±20, 576±34, and 583±35 g C m^?2 yr^?1 at HJP02, HJP94, HJP75, and OJP, respectively. The ratio of annual ecosystem respiration (R) to annual GEP was 2.51±0.15, 0.95±0.04, 0.87±0.03, and 0.96±0.03. Seasonally, NEP peaked in May or June at all four sites but GEP and R were highest in July. R at a reference soil temperature of 10 °C, ecosystem quantum yield and photosynthetic capacity were lowest for the 2‐year‐old stand. R was most sensitive to soil temperature for the 90‐year‐old stand. The primary source of variability in NEP over the course of succession of the jack pine ecosystem following harvesting was stand age due to the changes in leaf area index. Intersite variability in GEP and R was an order of magnitude greater than interannual variability at OJP. For both young and old stands, GEP had greater interannual variability than R and played a more important role than R in interannual variation in NEP. Based on year‐round flux measurements from 2000 to 2005, the 10‐year stand had larger interannual variability in GEP and R than the 90‐year stand. Interannual variability in NEP was driven primarily by early‐growing‐season temperature and growing‐season length. Photosynthesis played a dominant role in the rapid rise in NEP early in stand development. Late in stand development, however, the subtle decrease in NEP resulted primarily from increasing respiration.     

Role of vegetation in determining carbon sequestration along ecological succession in the southeastern United States

Vegetation plays a central role in controlling terrestrial carbon (C) exchange, but quantifying its impacts on C cycling on time scales of ecological succession is hindered by a lack of long‐term observations. The net ecosystem exchange of carbon (NEE) was measured for several years in adjacent ecosystems that represent distinct phases of ecological succession in the southeastern USA. The experiment was designed to isolate the role of vegetation – apart from climate and soils – in controlling biosphere–atmosphere fluxes of CO₂ and water vapor. NEE was near zero over 5 years at an early successional old‐field ecosystem (OF). However, mean annual NEE was nearly equal, approximately ?450 g C m^?2 yr^?1, at an early successional planted pine forest (PP) and a late successional hardwood forest (HW) due to the sensitivity of the former to drought and ice storm damage. We hypothesize that these observations can be explained by the relationships between gross ecosystem productivity (GEP), ecosystem respiration (RE) and canopy conductance, and long‐term shifts in ecosystem physiology in response to climate to maintain near‐constant ecosystem‐level water‐use efficiency (EWUE). Data support our hypotheses, but future research should examine if GEP and RE are causally related or merely controlled by similar drivers. At successional time scales, GEP and RE observations generally followed predictions from E. P. Odum's ‘Strategy of Ecosystem Development’, with the surprising exception that the relationship between GEP and RE resulted in large NEE at the late successional HW. A practical consequence of this research suggests that plantation forestry may confer no net benefit over the conservation of mature forests for C sequestration.     

The dependence of respiration on photosynthetic substrate supply and temperature: integrating leaf, soil and ecosystem measurements

Interactions between photosynthetic substrate supply and temperature in determining the rate of three respiration components (leaf, belowground and ecosystem respiration) were investigated within three environmentally controlled, Populus deltoides forest bays at Biosphere 2, Arizona. Over 2 months, the atmospheric CO₂ concentration and air temperature were manipulated to test the following hypotheses: (1) the responses of the three respiration components to changes in the rate of photosynthesis would differ both in speed and magnitude; (2) the temperature sensitivity of leaf and belowground respiration would increase in response to a rise in substrate availability; and, (3) at the ecosystem level, the ratio of respiration to photosynthesis would be conserved despite week‐to‐week changes in temperature. All three respiration rates responded to the CO₂ concentration‐induced changes in photosynthesis. However, the proportional change in the rate of leaf respiration was more than twice that of belowground respiration and, when photosynthesis was reduced, was also more rapid. The results suggest that aboveground respiration plays a key role in the overall response of ecosystem respiration to short‐term changes in canopy photosynthesis. The short‐term temperature sensitivity of leaf respiration, measured within a single night, was found to be affected more by developmental conditions than photosynthetic substrate availability, as the Q₁₀ was lower in leaves that developed at high CO₂, irrespective of substrate availability. However, the temperature sensitivity of belowground respiration, calculated between periods of differing air temperature, appeared to be positively correlated with photosynthetic substrate availability. At the ecosystem level, respiration and photosynthesis were positively correlated but the relationship was affected by temperature; for a given rate of daytime photosynthesis, the rate of respiration the following night was greater at 25 than 20°C. This result suggests that net ecosystem exchange did not acclimate to temperature changes lasting up to 3 weeks. Overall, the results of this study demonstrate that the three respiration terms differ in their dependence on photosynthesis and that, short‐ and medium‐term changes in temperature may affect net carbon storage in terrestrial ecosystems.     

腾格里荒漠红砂-珍珠群落CO2收支变化及其不同观测方法间的比较

由于荒漠生态系统植被覆盖度低、生产力低下,其在全球碳循环中的作用被长期忽视。为探讨荒漠生态系统碳收支各组分的变化规律,以腾格里荒漠红砂(Reaumuria soongorica Maxim.)-珍珠(Salsola passerina Beg.)群落为研究对象,采用静态箱式法研究了该群落的净生态系统CO2交换量(NEE)、生态系统呼吸、土壤呼吸的日变化规律,同时将该方法所获得的NEE结果与涡动相关法观测的结果进行了比较。结果表明:(1)红砂-珍珠群落NEE的日变化表现为,在6:00—9:00左右出现一个CO2吸收的高峰值,随后在12:00—15:00左右出现一个CO2释放高峰值。红砂种群、珍珠种群和整个群落NEE的平均值分别为0.018、0.020和0.028 mg CO2m-2s-1;(2)红砂种群、珍珠种群、土壤及整个群落生态系统呼吸速率的日变化规律一致,均表现为明显的单峰变化趋势,在12:00—15:00左右出现一个CO2释放的高峰值。红砂种群、珍珠种群、土壤和整个群落的生态系统呼吸的平均值分别为:0.121、0.062、0.029和0.040 mg CO2m-2s-1。以盖度为加权因子计算得到红砂种群、珍珠种群和土壤呼吸占生态系统呼吸的比例分别为:9%、21%和70%,由此可见,生态系统呼吸主要来源于土壤呼吸。(3)将箱式法和涡动相关法观测的NEE进行比较,结果表明两种方法观测的NEE变化规律基本一致,相关系数达到0.7。采用箱式法观测的NEE高于涡动相关法观测的结果,平均值分别0.028 mg CO2m-2s-1(箱式法)和0.015 mg CO2m-2s-1(涡动相关法),涡动相关法的观测结果与箱式法观测结果的比值为0.54。综上可得,荒漠生态系统土壤呼吸的变化速率决定了生态系统呼吸的变化规律,采用箱式法可能高估了荒漠生态系统CO2的释放量。     

Diurnal, seasonal and annual variation in net ecosystem CO2 exchange of an alpine shrubland on Qinghai-Tibetan plateau

Thus far, grassland ecosystem research has mainly been focused on low‐lying grassland areas, whereas research on high‐altitude grassland areas, especially on the carbon budget of remote areas like the Qinghai‐Tibetan plateau is insufficient. To address this issue, flux of CO₂ were measured over an alpine shrubland ecosystem (37°36′N, 101°18′E; 325 above sea level [a. s. l.]) on the Qinghai‐Tibetan Plateau, China, for 2 years (2003 and 2004) with the eddy covariance method. The vegetation is dominated by formation Potentilla fruticosa L. The soil is Mol–Cryic Cambisols. To interpret the biotic and abiotic factors that modulate CO₂ flux over the course of a year we decomposed net ecosystem CO₂ exchange (NEE) into its constituent components, and ecosystem respiration (R_eco). Results showed that seasonal trends of annual total biomass and NEE followed closely the change in leaf area index. Integrated NEE were ?58.5 and ?75.5 g C m^?2, respectively, for the 2003 and 2004 years. Carbon uptake was mainly attributed from June, July, August, and September of the growing season. In July, NEE reached seasonal peaks of similar magnitude (4–5 g C m^?2 day^?1) each of the 2 years. Also, the integrated night‐time NEE reached comparable peak values (1.5–2 g C m^?2 day^?1) in the 2 years of study. Despite the large difference in time between carbon uptake and release (carbon uptake time < release time), the alpine shrubland was carbon sink. This is probably because the ecosystem respiration at our site was confined significantly by low temperature and small biomass and large day/night temperature difference and usually soil moisture was not limiting factor for carbon uptake. In general, R_eco was an exponential function of soil temperature, but with season‐dependent values of Q₁₀. The temperature‐dependent respiration model failed immediately after rain events, when large pulses of R_eco were observed. Thus, for this alpine shrubland in Qinghai‐Tibetan plateau, the timing of rain events had more impact than the total amount of precipitation on ecosystem R_eco and NEE.     

Photosynthesis drives anomalies in net carbon-exchange of pine forests at different latitudes

The growth rate of atmospheric CO₂ exhibits large temporal variation that is largely determined by year‐to‐year fluctuations in land–atmosphere CO₂ fluxes. This land–atmosphere CO₂‐flux is driven by large‐scale biomass burning and variation in net ecosystem exchange (NEE). Between‐ and within years, NEE varies due to fluctuations in climate. Studies on climatic influences on inter‐ and intra‐annual variability in gross photosynthesis (GPP) and net carbon uptake in terrestrial ecosystems have shown conflicting results. These conflicts are in part related to differences in methodology and in part to the limited duration of some studies. Here, we introduce an observation‐driven methodology that provides insight into the dependence of anomalies in CO₂ fluxes on climatic conditions. The methodology was applied on fluxes from a boreal and two temperate pine forests. Annual anomalies in NEE were dominated by anomalies in GPP, which in turn were correlated with incident radiation and vapor pressure deficit (VPD). At all three sites positive anomalies in NEE (a reduced uptake or a stronger source than the daily sites specific long‐term average) were observed on summer days characterized by low incident radiation, low VPD and high precipitation. Negative anomalies in NEE occurred mainly on summer days characterized by blue skies and mild temperatures. Our study clearly highlighted the need to use weather patterns rather than single climatic variables to understand anomalous CO₂ fluxes. Temperature generally showed little direct effect on anomalies in NEE but became important when the mean daily air temperature exceeded 23 °C. On such days GPP decreased likely because VPD exceeded 2.0 kPa, inhibiting photosynthetic uptake. However, while GPP decreased, the high temperature stimulated respiration, resulting in positive anomalies in NEE. Climatic extremes in summer were more frequent and severe in the South than in the North, and had larger effects in the South because the criteria to inhibit photosynthesis are more often met.     

Model-data synthesis of diurnal and seasonal CO2 fluxes at Niwot Ridge, Colorado

Eddy covariance records hold great promise for understanding the processes controlling the net ecosystem exchange of CO₂ (NEE). However, NEE is the small difference between two large fluxes: photosynthesis and ecosystem respiration. Consequently, separating NEE into its component fluxes, and determining the process‐level controls over these fluxes, is a difficult problem. In this study, we used a model‐data synthesis approach with the Simplified PnET (SIPNET) flux model to extract process‐level information from 5 years of eddy covariance data at an evergreen forest in the Colorado Rocky Mountains. SIPNET runs at a twice‐daily time step, and has two vegetation carbon pools, a single aggregated soil carbon pool, and a soil moisture submodel that models both evaporation and transpiration. By optimizing the model parameters before evaluating model‐data mismatches, we were able to probe the model structure independent of any arbitrary parameter set. In doing so, we were able to learn about the primary controls over NEE in this ecosystem, and in particular the respiration component of NEE. We also used this parameter optimization, coupled with a formal model selection criterion, to investigate the effects of making hypothesis‐driven changes to the model structure. These experiments lent support to the hypotheses that (1) photosynthesis, and possibly foliar respiration, are down‐regulated when the soil is frozen and (2) the metabolic processes of soil microbes vary in the summer and winter, possibly because of the existence of distinct microbial communities at these two times. Finally, we found that including water vapor fluxes, in addition to carbon fluxes, in the parameter optimization did not yield significantly more information about the partitioning of NEE into gross photosynthesis and ecosystem respiration.     

Response of Net Ecosystem Productivity of Three Boreal Forest Stands to Drought

In 2001–03, continuous eddy covariance measurements of carbon dioxide (CO₂) flux were made above mature boreal aspen, black spruce, and jack pine forests in Saskatchewan, Canada, prior to and during a 3−year drought. During the 1st drought year, ecosystem respiration (R) was reduced at the aspen site due to the drying of surface soil layers. Gross ecosystem photosynthesis (GEP) increased as a result of a warm spring and a slow decrease of deep soil moisture. These conditions resulted in the highest annual net ecosystem productivity (NEP) in the 9 years of flux measurements at this site. During 2002 and 2003, a reduction of 6% and 34% in NEP, respectively, compared to 2000 was observed as the result of reductions in both R and GEP, indicating a conservative response to the drought. Although the drought affected most of western Canada, there was considerable spatial variability in summer rainfall over the 100−km extent of the study area; summer rainfalls in 2001 and 2002 at the two conifer sites minimized the impact of the drought. In 2003, however, precipitation was similarly low at all three sites. Due to low topographic position and consequent poor drainage at the black spruce site and the coarse soil with low water-holding capacity at the jack pine site almost no reduction in R, GEP, and NEP was observed at these two sites. This study shows that the impact of drought on carbon sequestration by boreal forest ecosystems strongly depends on rainfall distribution, soil characteristics, topography, and the presence of vegetation that is well adapted to these conditions.     

Warming impacts on boreal fen CO2 exchange under wet and dry conditions

Northern peatlands form a major soil carbon (C) stock. With climate change, peatland C mineralization is expected to increase, which in turn would accelerate climate change. A particularity of peatlands is the importance of soil aeration, which regulates peatland functioning and likely modulates the responses to warming climate. Our aim is to assess the impacts of warming on a southern boreal and a sub‐arctic sedge fen carbon dioxide (CO₂) exchange under two plausible water table regimes: wet and moderately dry. We focused this study on minerotrophic treeless sedge fens, as they are common peatland types at boreal and (sub)arctic areas, which are expected to face the highest rates of climate warming. In addition, fens are expected to respond to environmental changes faster than the nutrient poor bogs. Our study confirmed that CO₂ exchange is more strongly affected by drying than warming. Experimental water level draw‐down (WLD) significantly increased gross photosynthesis and ecosystem respiration. Warming alone had insignificant impacts on the CO₂ exchange components, but when combined with WLD it further increased ecosystem respiration. In the southern fen, CO₂ uptake decreased due to WLD, which was amplified by warming, while at northern fen it remained stable. As a conclusion, our results suggest that a very small difference in the WLD may be decisive, whether the C sink of a fen decreases, or whether the system is able to adapt within its regime and maintain its functions. Moreover, the water table has a role in determining how much the increased temperature impacts the CO₂ exchange.     

Forest soil CO2 flux: uncovering the contribution and environmental responses of ectomycorrhizas

Forests play a critical role in the global carbon cycle, being considered an important and continuing carbon sink. However, the response of carbon sequestration in forests to global climate change remains a major uncertainty, with a particularly poor understanding of the origins and environmental responses of soil CO₂ efflux. For example, despite their large biomass, the contribution of ectomycorrhizal (EM) fungi to forest soil CO₂ efflux and responses to changes in environmental drivers has, to date, not been quantified in the field. Their activity is often simplistically included in the ‘autotrophic’ root respiration term. We set up a multiplexed continuous soil respiration measurement system in a young Lodgepole pine forest, using a mycorrhizal mesh collar design, to monitor the three main soil CO₂ efflux components: root, extraradical mycorrhizal hyphal, and soil heterotrophic respiration. Mycorrhizal hyphal respiration increased during the first month after collar insertion and thereafter remained remarkably stable. During autumn the soil CO₂ flux components could be divided into ∼60% soil heterotrophic, ∼25% EM hyphal, and ∼15% root fluxes. Thus the extraradical EM mycelium can contribute substantially more to soil CO₂ flux than do roots. While EM hyphal respiration responded strongly to reductions in soil moisture and appeared to be highly dependent on assimilate supply, it did not responded directly to changes in soil temperature. It was mainly the soil heterotrophic flux component that caused the commonly observed exponential relationship with temperature. Our results strongly suggest that accurate modelling of soil respiration, particularly in forest ecosystems, needs to explicitly consider the mycorrhizal mycelium and its dynamic response to specific environmental factors. Moreover, we propose that in forest ecosystems the mycorrhizal CO₂ flux component represents an overflow ‘CO₂ tap’ through which surplus plant carbon may be returned directly to the atmosphere, thus limiting expected carbon sequestration from trees under elevated CO₂.     

Carbon sequestration and ecosystem respiration for 4 years in a Scots pine forest

The net exchange of CO₂ (NEE) between a Scots pine (Pinus sylvestris L.) forest ecosystem in eastern Finland and the atmosphere was measured continuously by the eddy covariance (EC) technique over 4 years (1999–2002). The annual temperature coefficient (Q₁₀) of ecosystem respiration (R) for these years, respectively, was 2.32, 2.66, 2.73 and 2.69. The light‐saturated rate of photosynthesis (A_max) was highest in July or August, with an annual average A_max of 10.9, 14.6, 15.3 and 17.1 μmol m^?2 s^?1 in the 4 years, respectively. There was obvious seasonality in NEE, R and gross primary production (GPP), exhibiting a similar pattern to photosynthetically active radiation (PAR) and air temperature. The integrated daily NEE ranged from 2.59 to ?4.97 g C m^?2 day^?1 in 1999, from 2.70 to ?4.72 in 2000, from 2.61 to ?4.71 in 2001 and from 5.27 to ?4.88 in 2002. The maximum net C uptake occurred in July, with the exception of 2000, when it was in June. The interannual variation in ecosystem C flux was pronounced. The length of the growing season, based on net C uptake, was 179, 170, 175 and 176 days in 1999–2002, respectively, and annual net C sequestration was 152, 101, 172 and 205 g C m^?2 yr^?1. It is estimated that ecosystem respiration contributed 615, 591, 752 and 879 g C m^?2 yr^?1 to the NEE in these years, leading to an annual GPP of ?768, ?692, ?924 and ?1084 g C m^?2 yr^?1. It is concluded that temperature and PAR were the main determinants of the ecosystem CO₂ flux. Interannual variations in net C sequestration are predominantly controlled by average air temperature and integrated radiation in spring and summer. Four years of EC data indicate that boreal Scots pine forest ecosystem in eastern Finland acts as a relatively powerful carbon sink. Carbon sequestration may benefit from warmer climatic conditions.     

Carbon sequestration in a high-elevation, subalpine forest

We studied net ecosystem CO₂ exchange (NEE) dynamics in a high‐elevation, subalpine forest in Colorado, USA, over a two‐year period. Annual carbon sequestration for the forest was 6.71 mol C m^?2 (80.5 g C m^?2) for the year between November 1, 1998 and October 31, 1999, and 4.80 mol C m^?2 (57.6 g C m^?2) for the year between November 1, 1999 and October 31, 2000. Despite its evergreen nature, the forest did not exhibit net CO₂ uptake during the winter, even during periods of favourable weather. The largest fraction of annual carbon sequestration occurred in the early growing‐season; during the first 30 days of both years. Reductions in the rate of carbon sequestration after the first 30 days were due to higher ecosystem respiration rates when mid‐summer moisture was adequate (as in the first year of the study) or lower mid‐day photosynthesis rates when mid‐summer moisture was not adequate (as in the second year of the study). The lower annual rate of carbon sequestration during the second year of the study was due to lower rates of CO₂ uptake during both the first 30 days of the growing season and the mid‐summer months. The reduction in CO₂ uptake during the first 30 days of the second year was due to an earlier‐than‐normal spring warm‐up, which caused snow melt during a period when air temperatures were lower and atmospheric vapour pressure deficits were higher, compared to the first 30 days of the first year. The reduction in CO₂ uptake during the mid‐summer of the second year was due to an extended drought, which was accompanied by reduced latent heat exchange and increased sensible heat exchange. Day‐to‐day variation in the daily integrated NEE during the summers of both years was high, and was correlated with frequent convective storm clouds and concomitant variation in the photosynthetic photon flux density (PPFD). Carbon sequestration rates were highest when some cloud cover was present, which tended to diffuse the photosynthetic photon flux, compared to periods with completely clear weather. The results of this study are in contrast to those of other studies that have reported increased annual NEE during years with earlier‐than‐normal spring warming. In the current study, the lower annual NEE during 2000, the year with the earlier spring warm‐up, was due to (1) coupling of the highest seasonal rates of carbon sequestration to the spring climate, rather than the summer climate as in other forest ecosystems that have been studied, and (2) delivery of snow melt water to the soil when the spring climate was cooler and the atmosphere drier than in years with a later spring warm‐up. Furthermore, the strong influence of mid‐summer precipitation on CO₂ uptake rates make it clear that water supplied by the spring snow melt is a seasonally limited resource, and summer rains are critical for sustaining high rates of annual carbon sequestration.     

Carbon exchange in biological soil crust communities under differential temperatures and soil water contents: implications for global change

Biological soil crusts (biocrusts) are an integral part of the soil system in arid regions worldwide, stabilizing soil surfaces, aiding vascular plant establishment, and are significant sources of ecosystem nitrogen and carbon. Hydration and temperature primarily control ecosystem CO₂ flux in these systems. Using constructed mesocosms for incubations under controlled laboratory conditions, we examined the effect of temperature (5–35 °C) and water content (WC, 20–100%) on CO₂ exchange in light (cyanobacterially dominated) and dark (cyanobacteria/lichen and moss dominated) biocrusts of the cool Colorado Plateau Desert in Utah and the hot Chihuahuan Desert in New Mexico. In light crusts from both Utah and New Mexico, net photosynthesis was highest at temperatures >30 °C. Net photosynthesis in light crusts from Utah was relatively insensitive to changes in soil moisture. In contrast, light crusts from New Mexico tended to exhibit higher rates of net photosynthesis at higher soil moisture. Dark crusts originating from both sites exhibited the greatest net photosynthesis at intermediate soil water content (40–60%). Declines in net photosynthesis were observed in dark crusts with crusts from Utah showing declines at temperatures >25 °C and those originating from New Mexico showing declines at temperatures >35 °C. Maximum net photosynthesis in all crust types from all locations were strongly influenced by offsets in the optimal temperature and water content for gross photosynthesis compared with dark respiration. Gross photosynthesis tended to be maximized at some intermediate value of temperature and water content and dark respiration tended to increase linearly. The results of this study suggest biocrusts are capable of CO₂ exchange under a wide range of conditions. However, significant changes in the magnitude of this exchange should be expected for the temperature and precipitation changes suggested by current climate models.     

Response of Net Ecosystem Productivity of Three Boreal Forest Stands to Drought

In 2000–03, continuous eddy covariance measurements of carbon dioxide (CO₂) flux were made above mature boreal aspen, black spruce, and jack pine forests in Saskatchewan, Canada, prior to and during a 3-year drought. During the 1st drought year, ecosystem respiration (R) was reduced at the aspen site due to the drying of surface soil layers. Gross ecosystem photosynthesis (GEP) increased as a result of a warm spring and a slow decrease of deep soil moisture. These conditions resulted in the highest annual net ecosystem productivity (NEP) in the 9 years of flux measurements at this site. During 2002 and 2003, a reduction of 6% and 34% in NEP, respectively, compared to 2000 was observed as the result of reductions in both R and GEP, indicating a conservative response to the drought. Although the drought affected most of western Canada, there was considerable spatial variability in summer rainfall over the 100-km extent of the study area; summer rainfalls in 2001 and 2002 at the two conifer sites minimized the impact of the drought. In 2003, however, precipitation was similarly low at all three sites. Due to low topographic position and consequent poor drainage at the black spruce site and the coarse soil with low water-holding capacity at the jack pine site almost no reduction in R, GEP, and NEP was observed at these two sites. This study shows that the impact of drought on carbon sequestration by boreal forest ecosystems strongly depends on rainfall distribution, soil characteristics, topography, and the presence of vegetation that is well adapted to these conditions. The online version of the original article can be found under doi:     

Stand age-related effects on soil respiration in a first rotation Sitka spruce chronosequence in central Ireland

The effect of stand age on soil respiration and its components was studied in a first rotation Sitka spruce chronosequence composed of 10‐, 15‐, 31‐, and 47‐year‐old stands established on wet mineral gley in central Ireland. For each stand age, three forest stands with similar characteristics of soil type and site preparation were used. There were no significant differences in total soil respiration among sites of the same age, except for the case of a 15‐year‐old stand that had lower soil respiration rates due to its higher productivity. Soil respiration initially decreased with stand age, but levelled out in the older stands. The youngest stands had significantly higher respiration rates than more mature sites. Annual soil respiration rates were modelled by means of temperature‐derived functions. The average Q ₁₀ value obtained treating all the stands together was 3.8. Annual soil respiration rates were 991, 686, 556, and 564 g C m^?2 for the 10‐, 15‐, 31‐, and 47‐year‐old stands, respectively. We used the trenching approach to separate soil respiration components. Heterotrophic respiration paralleled soil organic carbon dynamics over the chronosequence, decreasing with stand age to slightly increase in the oldest stand as a result of accumulated aboveground litter and root inputs. Root respiration showed a decreasing trend with stand age, which was explained by a decrease in fine root biomass over the chronosequence, but not by nitrogen concentration of fine roots. The decrease in the relative contribution of autotrophic respiration to total soil CO₂ efflux from 59.3% in the youngest stand to 49.7% in the oldest stand was explained by the higher activity of the root system in younger stands. Our results show that stand age should be considered if simple temperature‐based models to predict annual soil respiration in afforestation sites are to be used.