Article

Ecosystem respiration budget for a Pinus sylvestris stand in central Siberia

Authors:
  • V.N. Sukachev Institute of Forest
  • V.N. Sukachev Institute of forest Siberian branch of Russian Academy of Sciences, Russia, Krasnoyarsk
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Abstract

abstractUsing a ground-based and an above-canopy eddy covariance system in addition to stem respiration measurements, the annual respiratory fluxes attributable to soil, stems and foliage were determined for a Scots pine (Pinus sylvestris L.) forest growing in central Siberia. Night-time foliar respiration was estimated on the basis of the difference between fluxes measured below and above the canopy and the stem respiration measurements. Comparison of the effects of night-time turbulence on measured CO2 fluxes showed flux loss above the canopy at low wind speeds, but no such effect was observed for the ground-based eddy system. This suggests that problems with flow homogeneity or flux divergence (both of which would be expected to be greater above the canopy than below) were responsible for above-canopy losses under these conditions. After correcting for this, a strong seasonality in foliar respiration was observed. This was not solely attributable to temperature variations, with intrinsic foliar respiratory capacities being much greater in spring and autumn. The opposite pattern was observed for stem respiration, with the intrinsic respiratory capacity being lower from autumn through early spring. Maximum respiratory activity was observed in early summer. This was not simply associated with a response to higher temperatures but seemed closely linked with cambial activity and the development of new xylem elements. Soil respiration rates exhibited an apparent high sensitivity to temperature, with seasonal data implying a Q10 of about 7. We interpret this as reflecting covarying changes in soil microbial activity and soil temperatures throughout the snow-free season. Averaged over the two study years (1999 and 2000), the annual respiratory flux was estimated at 38.3 mol C m−2 a−1. Of this 0.61 was attributable to soil respiration, with stem respiration accounting for 0.21 and foliar respiration 0.18.

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... Noticeable peak periods of soil CO2 efflux are observed in the summer or early autumn whereas soil efflux is lowest during the often-long winters (e.g. Rayment and Jarvis 2000;Högberg et al. 2001;Shibistova et al. 2002a;Pumpanen et al. 2003a;Domisch et al. 2006). In addition to seasonal changes in temperature and moisture, the seasonal pattern of soil CO2 efflux is influenced by many factors; root production of boreal plants and as well as mycelial production of ectomycorrhizal fungi have been found to vary seasonally in northern ecosystems (Wallander et al. 1997;2001;Steinaker et al. 2010), which most likely also influence the temporal variation of forest soil CO2 efflux through root respiration and root-associated heterotrophic respiration. ...
... The peak CO2 efflux occurred in July-August as observed in many previous studies in boreal coniferous forests (e.g. Morén and Lindroth 2000;Högberg et al. 2001;Shibistova et al. 2002a;Domisch et al. 2006;Kolari et al. 2009). The highest soil CO2 efflux at 10°C was found in August as well and the lowest in May, similar to the temperature response pattern observed in a Siberian Scots pine forest (Shibistova et al. 2002a). ...
... Morén and Lindroth 2000;Högberg et al. 2001;Shibistova et al. 2002a;Domisch et al. 2006;Kolari et al. 2009). The highest soil CO2 efflux at 10°C was found in August as well and the lowest in May, similar to the temperature response pattern observed in a Siberian Scots pine forest (Shibistova et al. 2002a). The observed seasonality of temperature response in monthly models corresponded also well to the pattern reported for a temperate forest (Janssens and Pilegaard 2003), with greater Q10's and lower base respiration (i.e. ...
... We studied the seasonal pattern of soil CO 2 efflux rates using both eddy covariance and chamber techniques, also examining seasonal changes in microbial population structure and sources of spatial variability in soil CO 2 efflux rates. Data from this study form the basis of an accompanying paper, in which estimates of the annual respiratory balance of the stand at the ecosystem level are given (Shibistova et al., 2002). ...
... The inter-relationships between soil temperature and soil moisture in determining overall rates of soil CO 2 efflux are discussed further in Section 3.5. This lack of a fit of the model of Lloyd and Taylor (1994) is considered in much more detail in an accompanying paper with alternative formulations for describing the temperature dependence of soil respiration for this ecosystem also being given (Shibistova et al., 2002). A second possibility for the differences between the chamber and eddy covariance measurements might be that the eddy covariance system was systematically understimating fluxes due to suppressed or intermittent turbulence characteristics close to the forest floor (Janssens et al., 2001). ...
... Even when eddy covariance measurements are made above a canopy, there are sometimes indications that some " loss of flux " may occur for under conditions of low wind speeds at night (Goulden et al., 1996; Jarvis et al., 1997). We have examined this problem closely elsewhere (Shibistova et al., 2002) and conclude that, in contrast to the eddy covariance system installed above the forest, flux " losses " at low turbulence were not a problem for the ground eddy covariance system in this experiment. It is, however, well known that different soil CO 2 efflux measurement methodologies or even instruments can give vastly different soil CO 2 efflux estimates (Healy et al., 1996; Norman et al., 1997; Arneth et al., 1998; Le Dantec et al., 1999; Janssens et al., 2001), and the basis of these differences is not always well known or even consistent. ...
Article
Rates of CO 2 efflux from the floor of a central Siberian Scots pine (Pinus sylvestris) forest were measured using a dynamic closed chamber system and by a eddy covariance system placed 2.5 m above the forest floor. Measurements were undertaken for a full growing season: from early May to early October 1999. Spatial variability as determined by the chamber measurements showed the rate of CO 2 efflux to depend on location, with rates from relatively open areas ("glades") only being about 50% those observed below or around trees. This was despite generally higher temperatures in the glade during the day. A strong relationship between CO 2 efflux rate and root density was observed in early spring, suggesting that lower rates in open areas may have been attributable to fewer roots there. Continuous measurements with the eddy covariance system provided good temporal coverage. This method, however, provided estimates of ground CO 2 efflux rate rates that were about 50% lower than chamber measurements that were undertaken in areas considered to be representative of the forest as a whole. An examination of the seasonal pattern of soil CO 2 efflux rates suggests that much of the variability in CO 2 efflux rate could be accounted for by variations in soil temperature. Nevertheless, there were also some indications that the soil water deficits served to reduce soil CO 2 efflux rates during mid-summer. Overall the sensitivity of CO 2 efflux rate to temperature seems to be greater for this boreal ecosystem than has been the case for most other studies.
... Our interest lies with the underlying controls on canopy physiology, rather than reporting annual totals (for these c.f. above referenced publications), and we therefore divide the measured ecosystem-atmosphere exchange of CO 2 (NEE) into its component fluxes assimilation (A) and respiration (R). In case of the Siberian ecosystems, Arrhenius-type relationships of measured night-time NEE (=R) with soil temperature were established, and by using these, R were extrapolated to daylight hours (Lloyd and Taylor, 1994;Arneth et al., 2002a;Shibistova et al., 2002). In the Mopane woodland, ecosystem respiration rates were frequently confined by soil moisture rather than by temperature. ...
... This period thus exceeds the green or growing season, as it can commence before new leaf-growth and continue beyond senescence. We also explicitly acknowledge continuation of some ecosystem activity during the dormant period, particularly heterotrophic respiration, since the low rates of CO 2 efflux observed in the cold or dry season can sum up to a sizeable portion of the annual budget and must not be ignored (Hanan et al., 1998;Lafleur et al., 2001;Arneth et al., 2002a;Shibistova et al., 2002;Aurela et al., 2004;Veenendaal et al., 2004). To alleviate comparison of the seasonality in the observed carbon exchange rates in ecosystems that represent diverse biomes from both hemispheres we define the onset of the active period as the first day following the month of August when rain exceeds 10 mm (Botswana), and the first day after January with average air temperature exceeding 0 @BULLET C (Siberia), respectively. ...
... Still, separate cuvette measurements on soil, stem and leaf level would be required to investigate these processes in more detail. Throughout the year, an ecosystems' " capacity " for respiration varies since plant growth and maintenance respiration, and heterotrophic activity respond rather plastically to varying environmental conditions (e.g.,Law et al., 1999;Arneth et al., 2002a;Shibistova et al., 2002;Atkin and Tjoelker, 2003;Pendall et al., 2004). Over a period of weeks to months, the exponential respiration-temperature response which typically dominates the short-term variation in respiration rates is thus mediated by additional factors, like available soil moisture or phenology. ...
Article
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We compare assimilation and respiration rates, and water use strategies in four divergent ecosystems located in cold-continental central Siberia and in semi-arid southern Africa. These seemingly unrelated systems have in common a harsh and highly seasonal environment with a very sharp transition between the dormant and the active season, and with vegetation facing dry air and soil conditions for at least part of the year. Moreover, the northern high latitudes and the semi-arid tropics will likely experience changes in key environmental parameters (e.g., air temperature and precipitation) in the future; indeed, in some regions marked climate trends have already been observed over the last decade or so. The magnitude of instantaneous or daily assimilation and respiration rates, derived from one to two years of eddy covariance measurements in each of the four ecosystems, was not related to the growth environment. For instance, respiration rates were clearly highest in the two deciduous systems included in the analysis (a Mopane woodland in northern Botswana and a Downy birch forest in Siberia; > 300 mmol m-2Td-1), while assimilation rates in the Mopane woodland were relatively similar to a Siberian Scots pine canopy for a large part of the active season (ca. 420 mmol m -2d-1). Acknowledging the limited number of ecosystems compared here, these data nevertheless suggest that factors like vegetation type, canopy phenology or ecosystem age can override larger-scale climate differences in terms of their effects on carbon assimilation and respiration rates. By far the highest rates of assimilation were observed in Downy birch, an early successional species. These were achieved at a rather conservative water use, as indicated by relatively low levels of A, the marginal water cost of plant carbon gain. Surprisingly, the Mopane woodland growing in the semi-arid environment had significantly higher values of ¿. However, its water use strategy included a very plastic response to intermittently dry periods, and values of A were much more conservative overall during a rainy season with low precipitation and high air saturation deficits. Our comparison demonstrates that forest ecosystems can respond very dynamically in terms of water use strategy, both on interannual and much shorter time scales. It remains to be evaluated whether and in which ecosystems this plasticity is mainly due to a short-term stomatal response, or goes hand in hand with changes in canopy photosynthetic capacity
... The contribution of forest floor respiration to the total ecosystem respiration observed in our study was close to the higher end of the range of 40-90% estimated for other boreal forests (Bergeron et al., 2009;Black et al., 1996;Goulden and Crill, 1997;Ikawa et al., 2015;Launiainen et al., 2005;Morén and Lindroth, 2000;Shibistova et al., 2002). The stem and foliage respiration estimated as R eco minus R ff , accounting for 10-32% of R eco at our pine stand, compares with 39% of R eco estimated in a 200-year-old Scots pine stand in central Siberia (Shibistova et al., 2002) and 15-45% of R eco reported in a temperate 70-year-old white pine (Pinus strobues L.) forest in Canada (Peichl et al., 2010). ...
... The contribution of forest floor respiration to the total ecosystem respiration observed in our study was close to the higher end of the range of 40-90% estimated for other boreal forests (Bergeron et al., 2009;Black et al., 1996;Goulden and Crill, 1997;Ikawa et al., 2015;Launiainen et al., 2005;Morén and Lindroth, 2000;Shibistova et al., 2002). The stem and foliage respiration estimated as R eco minus R ff , accounting for 10-32% of R eco at our pine stand, compares with 39% of R eco estimated in a 200-year-old Scots pine stand in central Siberia (Shibistova et al., 2002) and 15-45% of R eco reported in a temperate 70-year-old white pine (Pinus strobues L.) forest in Canada (Peichl et al., 2010). However, the stem and foliage respiration might be slightly underestimated in this study because R ff measured using the EC method includes small fractions of the autotrophic respiration from stems, branches, foliage, and seedlings existing in the lower part of the forest below the EC mounting height (2.5 m), which is however likely small compared to the forest floor contribution (Tarvainen et al., 2017). ...
Article
The forest floor provides an important interface of soil-atmosphere CO2 exchanges but their controls and contributions to the ecosystem-scale carbon budget are uncertain due to measurement limitations. In this study, we deployed eddy covariance systems below- and above-canopy to measure the spatially integrated net forest floor CO2 exchange (NFFE) and the entire net ecosystem CO2 exchange (NEE) at two mature contrasting stands located in close vicinity in boreal Sweden. We first developed an improved cospectra model to correct below-canopy flux data. Our empirical below-canopy cospectra models revealed a greater contribution of large- and small-scale eddies in the trunk space compared to their distribution in the above-canopy turbulence cospectra. We found that applying the above-canopy cospectra model did not affect the below-canopy annual CO2 fluxes at the sparse pine forest but significantly underestimated fluxes at the dense mixed spruce-pine stand. At the mixed spruce-pine stand, forest floor respiration (Rff) was higher and photosynthesis (GPPff) was lower, leading to a 1.4 times stronger net CO2 source compared to the pine stand. We further found that drought enhanced Rff more than GPPff, leading to increased NFFE. Averaged across the six site-years, forest floor fluxes contributed 82% to ecosystem-scale respiration (Reco) and 12% to gross primary production (GPP). Since the annual GPP was similar between both stands, the considerable difference in their annual NEE was due to contrasting Reco, the latter being primarily driven by the variations in NFFE. This implies that NFFE acted as the driver for the differences in NEE between these two contrasting stands. This study therefore highlights the important role of forest floor CO2 fluxes in regulating the boreal forest carbon balance. It further calls for extended efforts in acquiring high spatiotemporal resolution data of forest floor fluxes to improve predictions of global change impacts on the forest carbon cycle.
... Nonetheless, several studies suggest the importance of starch accumulation in forest ecosystems, and this may account for discrepancies between interannual variations in eddy-covariance-based and biometricbased estimates. For example, interannual changes in tower-based GPP estimates did not correlate with changes in tree rings in a Scots pine forest growing in central Siberia (Shibistova et al. 2002). These data suggested that interannual changes in the demand of carbohydrates for new stem production might be compensated by changes in growth rates of other parts of the tree or by increased starch accumulation. ...
Chapter
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Net primary production (NPP) refers to the net amount of the carbon and energy fixed by green plants through photosynthetic activity. Estimates of NPP are of fundamental human importance, because food supply is predominantly dependent on plant productivity. Moreover, measurements of spatiotemporal variations of forest NPP provide important information for understanding and projecting the global carbon cycle, because forest ecosystems are a major terrestrial carbon sink. Here we discuss methods for estimating NPP in forest ecosystems using inventory data, and describe the “summation method”, which was developed in Japan in the 1960s at the International Biological Program (IBP) to facilitate standardization in the absence of complex instruments under field conditions. Global climate change prompted to development of this “summation method” as an improved “biometric method” in the 1990s. Biometric-based estimates of NPP are conceptually defined as the total amount of new organic matter produced during an interval per unit area at the ecosystem scale, and are expressed as the sum of stand increments of living biomass (SI), newly produced aboveground litter (L an ), and fine root production (P fr ). The SI of above- and belowground (coarse roots) biomass can be estimated by tracking the survival and diameter of individual tree stems in a permanent plot. Aboveground litter of short-lived organs (L an ), such as deciduous leaves, flowers, and fruits, can be determined using litter traps that are set on the forest floor. Although methods for determining P fr remain unstandardized, 1-year turnover of fine roots is often considered an estimate of fine root dynamics (P fr ≈ mean fine root biomass). In a study of the Takayama Experimental Forest, we demonstrated correlations of biometric-based NPP estimates with various methods, such as simulated canopy photosynthesis by scaling up leaf photosynthesis and incorporating values obtained using eddy covariance. The resulting biometric method has sufficient sensitivity to demonstrate climate-induced year-to-year variations of tree growth and allocation of carbon inputs by NPP in forest ecosystems.
... Correspondingly, in the Russian Federation, this method is constantly, temporarily, or occasionally used only in some sites, mainly in boreal forests. These studies were organized in the Central Forest Reserve, Tver oblast (Ol'chev et al., 2008); Krasnoyarsk krai, Zotino and Khakassia (Kelliher et al., 1998; Knohl et al., 2002; Shibistova et al., 2002); Central and Northern Yakutia (Maksii mov et al., 2005); ...
Article
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A lot of studies on the impact of global climate changes on natural communities deal with cryogenic ecosystems, tundra in particular, since they are delimited by low air temperature and permafrost, thus being extremely sensitive to long-term climate fluctuations. Continuous warming in Northern Hemisphere is unmasking all the more details concerning complex system of direct relationships, feedbacks, and interactions of carbon balance factors as the main response function. While the set of such factors may be viewed as more or less complete, their relative contribution to C-balance, as is becoming clear with accumulating results of field observations, directly depends on temporal scale of observations and is not constant. As the results of field observations and modeling of tundra ecosystems show, any one of significant factors can become the leading one within the boundaries determined by the given scale of observations. Even the least significant factor can become the determining one for direction of carbon annual net flux in an ecosystem, if contributions of more significant factors canceled each other during the period of observations. In the most general situation, the greater is the variation of a significant factor during the period of observations, the larger is its partial contribution. The complete set of independent variables of C-balance is not limited by abiotic factors but should include such an important factor as a stock of plants living top mass, which can be treated as not only the natural product of C-balance but also as its independent parameter.
... Moreover, chambers can change the soil-plant environment (air temperature and humidity) and affect the measurement results. For high precision and the ability to measure plant carbon flux at different heights, the eddy covariance technique is the main tool used to measure carbon flux in many FLUXNET sites (Valentini et al. 2000; Baldocchi 2003; giasson et al. 2006). in eddy covariance research, daytime ecosystem respiration is obtained from the empirical function of night time Co 2 exchange versus soil temperature (the Q 10 model) (Shibistova et al. 2002; Zamolodchikov et al. 2003; Zha et al. 2004; Xu & Baldocchi 2004; Van Dijk & Dolman 2004). This method is suitable for simulating ecosystem respiration of forest, grassland and single cropping systems. ...
Article
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CO2 flux was measured continuously using the eddy covariance technique in a wheat‐maize rotation system in the North China Plains from October 2002 to October 2006. The annual and seasonal variation of ecosystem respiration and the bio‐environmental controls on them were investigated. The results show that ecosystem respiration (Rec) in the cropland increased exponentially with soil temperature at 5 cm depth. The temperature sensitivity coefficient (Q10) for ecosystem respiration varied from 3.5 to 5.4 for wheat and from 2.4 to 4.5 for maize. In the wheat growing season, monthly average R0 (ecosystem respiration at 0°C) increased linearly with soil temperature and logarithmically with leaf area index (LAI). Monthly average Q10 decreased logarithmically with R0. Residual Rec was significantly correlated with LAI. After considering LAI, the modified Q10 model could estimate Rec better than before. The simulation results show that annual ecosystem respiration in the wheat‐maize rotation system in the North China Plains was 1327, 1348, 1040 and 1171 gC m yr for the 4 years of the study. As a 4‐year average, seasonal mean ecosystem respiration in wheat (2.60 gC m day) was much lower than in maize (6.09 gC m day). However, integrated ecosystem respiration for the wheat growing season (566 gC m) was slightly higher than that for maize (520 gC m). These account for 46.4 and 42.6% of the annual values, respectively.
... Monthly averages and models agreed with earlier findings of greater efflux in autumn compared to spring at comparable temperatures in a temperate forest (Crill, 1991). Our monthly models resulted in a pattern of predicted soil CO 2 efflux identical to that found in a Siberian pine forest study (Shibistova et al., 2002), with the month of May having the lowest CO 2 efflux at 10 @BULLET C and August having the highest. Variation in soil moisture did not explain the seasonality of the temperature response. ...
Article
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Our objectives were to identify factors related to temporal variation of soil CO2 efflux in a boreal pine forest and to evaluate simple predictive models of temporal variation of soil CO2 efflux. Soil CO2 efflux was measured with a portable chamber in a Finnish Scots pine forest for three years, with a fourth year for model evaluation. Plot averages for soil CO2 efflux ranged from 0.04 to 0.90 gCO2 m−2 h−1 during the snow-free period, i.e. May–October, and from 0.04 to 0.13 gCO2 m−2 h−1 in winter. Soil temperature was a good predictor of soil CO2 efflux. A quadratic model of lntransformed efflux explained 76–82% of the variation over the snow-free period. The results revealed an effect of season: at a given temperature of the organic layer, soil CO2 efflux was higher later in the snow-free period (in August and September) than in spring and early summer (in May and June). Regression coefficients for temperature (approximations of a Q10 value) of month-specific models decreased with increasing average soil temperatures. Efflux in July, the month of peak photosynthesis, showed no clear response to temperature or moisture. Inclusion of a seasonality index, degree days, improved the accuracy of temperature response models to predict efflux for the fourth year of measurements, which was not used in building of regression models. During peak efflux from mid-July to late-August, efflux was underestimated with the models that included degree days as well as with the models that did not. The strong influence of the flux of photosynthates belowground and the importance of root respiration could explain the relative temperature insensitivity observed in July and together with seasonality of growth of root and root-associated mycorrhizal fungi could explain partial failure of models to predict magnitude of efflux in the peak season from mid-July to August. http://www.biogeosciences.net/8/3169/2011/bg-8-3169-2011.pdf
... data reported by Quegan et al. ( 2011 ) – between 73 and 59 % of NPP. Similarly, Wirth et al. ( 2002 ) estimated 48 % for post fi re chronosequences in pine forests of Siberian middle taiga, although reported values for NEP (or NEE) for taiga forests of Central Siberia vary greatly – from 50–60 to 250–270 gC m −2 year −1 (e.g., Röser et al . 2002 ; Shibistova et al . 2002 ; Schulze 2002 ) . Substantial differences between NPP and soil respiration can occur, particularly for crops, since a major part of the NPP is removed as yield, e.g., the sink of agro-ecosystems of the taiga zone of the region was estimated to be 111 gC m −2 year −1 , or 42 % of NPP (Vedrova 2002 ) . Such effects usually are not represen ...
Chapter
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This chapter considers the current state of Siberian terrestrial ecosystems, their spatial distribution, and major biometric characteristics. Ongoing climate change and the dramatic increase of accompanying anthropogenic pressure provide different but mostly negative impacts on Siberian ecosystems. Future climates of the region may lead to substantial drying on large territories, acceleration of disturbance regimes, deterioration of ecosystems, and positive feedback to global warming. The region requires urgent development and implementation of strategies of adaptation to, and mitigation of, negative consequences of climate change.
... As the flux of melt water increases, the processes of freezing and thawing maintain a constant temperature near 0 1C in the upper soil layer. This phenomenon has been observed over extended periods mostly during autumn freeze-back in permafrost areas (Outcalt et al., 1990; Boike et al., 1998; Hinkel et al., 2001; Kane et al., 2001 ), and has also been previously observed throughout entire winters (Milyukova et al., 2002) or for extended periods after the first snowmelt in nonpermafrost boreal forest areas (Shibistova et al., 2002b). In our study, a zero-curtain period of 2–4 weeks preceded soil thaw in most springs with sub-zero soil temperatures. ...
Article
The timing of the commencement of photosynthesis (P*) in spring is an important determinant of growing-season length and thus of the productivity of boreal forests. Although controlled experiments have shed light on environmental mechanisms triggering release from photoinhibition after winter, quantitative research for trees growing naturally in the field is scarce. In this study, we investigated the environmental cues initiating the spring recovery of boreal coniferous forest ecosystems under field conditions. We used meteorological data and above-canopy eddy covariance measurements of the net ecosystem CO2 exchange (NEE) from five field stations located in northern and southern Finland, northern and southern Sweden, and central Siberia. The within- and intersite variability for P* was large, 30–60 days. Of the different climate variables examined, air temperature emerged as the best predictor for P* in spring. We also found that ‘soil thaw’, defined as the time when near-surface soil temperature rapidly increases above 0°C, is not a useful criterion for P*. In one case, photosynthesis commenced 1.5 months before soil temperatures increased significantly above 0°C. At most sites, we were able to determine a threshold for air-temperature-related variables, the exceeding of which was required for P*. A 5-day running-average temperature (T5) produced the best predictions, but a developmental-stage model (S) utilizing a modified temperature sum concept also worked well. But for both T5 and S, the threshold values varied from site to site, perhaps reflecting genetic differences among the stands or climate-induced differences in the physiological state of trees in late winter/early spring. Only at the warmest site, in southern Sweden, could we obtain no threshold values for T5 or S that could predict P* reliably. This suggests that although air temperature appears to be a good predictor for P* at high latitudes, there may be no unifying ecophysiological relationship applicable across the entire boreal zone.
... The final two scenarios have been taken in order to reflect a typical situation for daytime canopy photosynthesis and for a nighttime ecosystem respectively. Nocturnal respiration of forest ecosystems is typically dominated by the soil CO 2 efflux (Law et al., 1999; Janssens et al., 2001; Milyukova et al., 2002; Shibistova et al., 2002a). As can be seen inFigure 4B these different assumptions regarding the source distribution lead to significantly different footprint estimates. ...
Article
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A knowledge of the distribution of the contribution of upwind sources to measurements of vertical scalar flux densities is important for the correct interpretation of eddy covariance data. Several approaches have been developed to estimate this so-called footprint function. Here a new approach based on the ensemble-averaged Navier—Stokes equations is presented. Comparisons of numerical results using this approach with results from other studies under a range of environmental conditions show that the model predictions are robust. Moreover, the approach outlined here has the advantage of a potential wide applicability, due to an ability to take into account the heterogeneous nature of underlying surfaces. For example, the model showed that any variations in surface drag, such as must occur in real life heterogeneous canopies, can exert a marked influence of the shape and extent of flux footprints. Indeed, it seems likely that under such circumstances, estimates of surface fluxes will be weighted towards areas of highest foliage density (and therefore quite likely higher photosynthetic rates) close to the measurement sensor. Three-dimensional footprints during the day and night were also determined for a mixed coniferous forest in european Russia. A marked asymmetry of the footprint in the crosswind direction was observed, this being especially pronounced for non-uniform plant distributions involving vegetation types with different morphological and physiological properties. The model also found that, other things being equal, the footprint peak for forest soil respiration is typically over twice the distance from the above canopy measurement sensor compared to that for canopy photosynthesis. This result has important consequences for the interpretation of annual ecosystem carbon balances by the eddy covariance method.
... change in CO 2 concentration in the canopy airspace between the two devices (DC store is provided by gas profile measurements). The trunk respiration R st is estimated at 0.3 and 0.4 mmol m À2 s À1 for 1999 and 2000, respectively [Shibistova et al., 2002]. We follow previous authors in designating skyward carbon flow as positive, although, when quantifying photosynthesis and respiration, we discuss them in terms of their magnitude. ...
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This paper was published as Journal of Geophysical Research, 2005, 110, D23209. Copyright © 2005 American Geophysical Union. It is also available from the publisher's website at http://www.agu.org/pubs/crossref/2005/2005JD006060.shtml. Doi: 10.1029/2005JD006060 We have expanded the Monte Carlo, ray-tracing model FLIGHT in order to simulate photosynthesis within three-dimensional, heterogeneous tree canopies. In contrast to the simple radiative transfer schemes adopted in many land-surface models (e.g., the Big Leaf approximation), our simulation calculates explicitly the leaf irradiance at different heights within the canopy and thus produces an accurate scale-up in photosynthesis from leaf to canopy level. We also account for both diffuse and direct sunlight. For a Siberian stand of Scots pine Pinus sylvestris, FLIGHT predicts observed carbon assimilation, across the full range of sky radiance, with an r.m.s. error of 12%. Our main findings for this sparse canopy, using both measurements and model, are as follows: (1) Observationally, we detect a light-use efficiency (LUE) increase of only ≤10% for the canopy when the proportion of diffuse sky radiance is 75% rather than 25%. The corresponding enhancement predicted by our simulations is 10–20%. With such small increases in LUE, our site will not assimilate more carbon on overcast days compared to seasonally equivalent sunny days; (2) the scale-up in photosynthesis from top-leaf to canopy is less than unity. The Big Leaf approximation, based on Beer's law and light-acclimated leaf nitrogen, overpredicts this scale-up by ≥60% for low sky radiance (≤500 μmolPAR m−2 s−1); (3) when leaf nitrogen is distributed so as to maximize canopy photosynthesis, the increase in the canopy carbon assimilation, compared with a uniform nitrogen distribution, is small (≅4%). Maximum assimilation occurs when the vertical gradient of leaf nitrogen is slightly shallower than that of the light profile.
... [13] Several studies were previously conducted close to the ZOTTO site. Eddy covariance flux measurements of CO 2 , H 2 O, and energy exchanges were made in a nearby pine forest Shibistova et al., 2002b], dark taiga [Röser et al., 2002], and bogs [Kurbatova et al., 2002]. These measurements were accompanied by process studies on soil respiration [Shibistova et al., 2002a] and detailed forest inventories Wirth et al., 1999]. ...
Article
We present first results from 19 months of semicontinuous concentration measurements of biogeochemical trace gases (CO2, CO, and CH4) and O2, measured at the Zotino Tall Tower Observatory (ZOTTO) in the boreal forest of central Siberia. We estimated CO2 and O2 seasonal cycle amplitudes of 26.6 ppm and 134 per meg, respectively. An observed west-east gradient of about −7 ppm (in July 2006) between Shetland Islands, Scotland, and ZOTTO reflects summertime continental uptake of CO2 and is consistent with regional modeling studies. We found the oceanic component of the O2 seasonal amplitude (Atmospheric Potential Oxygen, or APO) to be 51 per meg, significantly smaller than the 95 per meg observed at Shetlands, illustrating a strong attenuation of the oceanic O2 signal in the continental interior. Comparison with the Tracer Model 3 (TM3) atmospheric transport model showed good agreement with the observed phasing and seasonal amplitude in CO2; however, the model exhibited greater O2 (43 per meg, 32%) and smaller APO (9 per meg, 18%) amplitudes. This seeming inconsistency in model comparisons between O2 and APO appears to be the result of phasing differences in land and ocean signals observed at ZOTTO, where ocean signals have a significant lag. In the first 2 months of measurements on the fully constructed tower (November and December 2006), we observed several events with clear vertical concentration gradients in all measured species except CO. During “cold events” (below −30°C) in November 2006, we observed large vertical gradients in CO2 (up to 22 ppm), suggesting a strong local source. The same pattern was observed in CH4 concentrations for the same events. Diurnal vertical CO2 gradients in April to May 2007 gave estimates for average nighttime respiration fluxes of 0.04 ± 0.02 mol C m−2 d−1, consistent with earlier eddy covariance measurements in 1999–2000 in the vicinity of the tower.
... The observed 4-week period of constant near-zero temperatures during snow melting in the spring—the zero curtain—is a manifestation of the temperature-balancing effect of latent heat absorption by melting water and release by freezing water until all the water in the soil has melted or frozen completely. This behavior has been reported in permafrost areas mostly during autumn (Outcalt et al. 1990, Boike et al. 1998) and also throughout entire winters (Milyukova et al. 2002) or after the first snowmelt in non-permafrost boreal forest areas (Shibistova et al. 2002 ), but has not been observed at our site before , probably because the soil has never before frozen so completely during the monitoring period. Suni et al. (2003a) reported a zero-curtain effect at several boreal sites after wintertime soil temperatures several degrees below zero, but never when wintertime temperatures remained close to or above zero. ...
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... ld then be sealed to the body of a container mounted on a tree stem by two locking clips. The bodies of the containers were sealed to tree stems with a silicon sealant, and remained in place between stem CO 2 efflux measurements. We checked the chambers for leaks by exhaling near the seals and monitoring the CO 2 concentration inside the chambers (Shibistova et al . 2002). The chamber volumes were determined following the method of Edwards & Hanson (1996); the mean chamber volume was 793 ± 20 mL (mean ± 1 SD). The mean total system volume including the infrared gas analyser (IRGA) was 958 ± 21 mL (mean ± 1 SD). Each chamber covered an area of 150 cm 2 of stem surface. Thermocouples were inserted to appr ...
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Mean residence time (MRT) of topsoil organic carbon is one critical parameter for predicting future land carbon sink dynamics. Large uncertainties remain about controls on the variability in global MRT of soil organic carbon. We estimated global MRT of topsoil (0-20 cm) organic carbon in terrestrial ecosystems and found that mean annual air temperature, annual precipitation, and topsoil nitrogen storage were responsible for the variability in MRT. An empirical climate and soil nitrogen - based (Clim&SN) model could be used to explain the temporal and spatial variability in MRT across various ecosystems. Estimated MRT was lowest in the low-latitude zones, and increased toward high-latitude zones. Global MRT of topsoil organic carbon showed a significant declining tendency between 1960 and 2008, particularly in the high-latitude zone of the northern hemisphere. The largest absolute and relative changes (0.2% per yr) in MRT of topsoil organic carbon from 1960 to 2008 occurred in high-latitude regions, consistent with large carbon stocks in, and greater degree of climate change being experienced by, these areas. Overall, global MRT anomalies (differences between MRT in each year and averaged value of MRT from 1960 to 2008) of terrestrial topsoil organic carbon were decreasing from 1960 to 2008. Global MRT anomalies decreased significantly (P<0.001) with the increase of global temperature anomalies, indicating that global warming resulted in faster turnover rates of topsoil organic carbon.
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Tower‐based eddy covariance measurements were used to quantify the effect of fire on subsequent carbon dioxide fluxes and water and surface energy balance characteristics for campo sujo savanna located near Brasília in Central Brazil (15°56′ S, 47°51′ W). Campo sujo is a xeromorphic, open shrub savanna with very scattered but definitely visible shrubs and tree‐like shrub elements. We studied two areas, one exposed to a prescribed fire late in the dry season, and a second that had not been burned for the previous 4 years. The fire on 22 September 1998 consumed an estimated 26 mol C m ⁻² . Immediately after the fire, evapotranspiration rates decreased and the savanna became a stronger net source of CO 2 to the atmosphere. This was attributed to the removal of the still slightly physiologically active grass layer and higher soil CO 2 efflux rates as a consequence of elevated surface soil temperatures post‐burning. On the commencement of the first rains in early October 1998, this situation was reversed, with the burned area rapidly becoming a stronger sink for CO 2 and with higher evapotranspiration rates than a nearby unburned (control) area. This difference persisted throughout the wet season (until at least June 1999) and was attributable to greater physiological activity of the regrowing vegetation in the burned area. Early in the growing season, higher soil evaporation rates may also have contributed to faster water use by the previously burned area. Overall, we estimate an annual gross primary productivity for the burned area of 135 mol C m ⁻² year ⁻¹ , with that for the unburned area being 106 mol C m ⁻² year ⁻¹ . Estimated ecosystem respiration rates were more similar on an annual basis (96 and 82 mol C m ⁻² year ⁻¹ for the burned and unburned areas, respectively), giving rise to a substantially higher net ecosystem productivity for the previously burned area (38 vs 24 mol C m ⁻² year ⁻¹ ). Stimulation of photosynthetic activity in the rapid post‐fire growth phase means that the negative effects of fire on the ecosystem carbon balance were more or less neutralized after only 12 months.
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Our objectives were to identify factors related to temporal variation of soil CO2 efflux in a boreal pine forest and to evaluate simple predictive models of temporal variation of soil CO2 efflux. Soil CO2 efflux was measured with a portable chamber in a Finnish Scots pine forest for three years, with a fourth year for model evaluation. Plot averages for soil CO2 efflux ranged from 0.04 to 0.90 g CO2 m-2 h-1 during the snow-free period, i.e. May-October, and from 0.04 to 0.13 g CO2 m-2 h-1 in winter. Soil temperature was a good predictor of soil CO2 efflux. A quadratic model of ln-transformed efflux explained 76-82% of the variation over the snow-free period. The results revealed strong seasonality: at a given soil temperature, soil CO2 efflux was higher later in the snow-free period than in spring and early summer. Regression coefficients for temperature (approximations of a Q10 value) of month-specific models decreased with increasing average soil temperatures. Efflux in July, the month of peak photosynthesis, showed no clear response to temperature or moisture. Inclusion of a seasonality index, degree days, improved the accuracy of temperature response models to predict efflux for the fourth year of measurements, which was not used in building of regression models. Underestimation during peak efflux (mid-July to late-August) remained uncorrected. The strong influence of the flux of photosynthates belowground and the importance of root respiration could explain the relative temperature insensitivity observed in July and together with seasonality of growth of root and root-associated mycorrhizal fungi could explain partial failure of models to predict magnitude of efflux in the peak season from mid-July to August. The effect of moisture early in the season was confounded by simultaneous advancement of the growing season and increase in temperature. In a dry year, however, the effect of drought was evident as soil CO2 efflux was some 30% smaller in September than in the previous wet year. Although soil temperature was a good overall predictor of soil CO2 efflux, possibly partly due to its proxy-like quality for covarying processes, strong seasonality of the temperature response observed in this boreal forest corroborates recent findings concerning the importance of seasonal changes in carbon inputs to processes producing CO2 in soil.
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Annual measurements of the diameter growth and litter fall of trees began in 1998 using a 1.0 ha permanent plot beneath a flux tower at the Takayama flux site, central Japan. This opened up an opportunity for studies that compare the interannual variability in tree growth with eddy covariance-based net ecosystem production (NEP). A possible link between multiyear biometric-based net primary production (NPP) and eddy covariance-based NEP was investigated to determine the contribution of autotrophic production and heterotrophic respiration (HR) to the interannual variability of NEP in deciduous forest ecosystems. We also defined the NEP* as the measurable organic matter stored in an ecosystem during the interval in which soil respiration (SR) measurements were taken. The difference of biometric-based NEP* from eddy covariance-based NEP within a given year varied between 55% and 105%. Woody tissue NPP (stems and coarse roots) varied markedly from 0.88 to 1.96 Mg C ha−1 yr−1 during the 8-year study period (1999–2006). Annual woody tissue NPP was positively correlated with eddy covariance-based NEP (r2=0.52, P<0.05). However, neither foliage NPP (r2=0.03) nor HR (r2=0.06) were correlated with eddy covariance-based NEP. Therefore, it was hypothesized that interannual variability in the ecosystem carbon exchange was directly responsible for much of the interannual variation in autotrophic production, especially carbon accumulation in the woody components of the ecosystem. Moreover, similar interannual variations of biometric-based NEP* and eddy covariance-based NEP with small variations in SR and foliage NPP suggest a constant net accumulation of carbon in nonliving pools at the Takayama site.
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Northern Eurasia is the largest terrestrial reservoir of carbon, and its dynamics and interactions with climate are globally significant. We present five independent estimates of the contemporary carbon balance of central Siberia using three different methodologies: a landscape-ecosystem approach (LEA) that amalgamates comprehensive vegetation, soil, hydrological and morphological information into a Geographical Information System, linked to regression-based estimates of carbon flux; two Dynamic Global Vegetation Models (DGVMs); and two atmospheric inversions. Apart from one of the DGVMs, all methods produce estimates of the net biome productivity (NBP) that are consistent both amongst themselves and with a range of other estimates. They indicate the region to be a carbon sink with a NBP of 27.5 ± 7.2 g C m−2 yr−1, which is equivalent to 352 ± 92 Mt C yr−1 if considered representative for boreal Asia. This is comparable with fossil fuel emissions for the Russian Federation, currently estimated as 427 MtC yr−1, and implies that boreal Asia does not play the major role in the northern hemisphere land sink, typically estimated to be of magnitude 1.5–2.9 Gt C yr−1. The LEA and DGVM approaches produce very different partitioning of NBP into its component fluxes. The DGVMs find net primary production (NPP) to be nearly balanced by heterotrophic respiration, disturbance being a relatively small term pushing the system closer to equilibrium. In the LEA, heterotrophic respiration is significantly less than NPP, and disturbance plays a much larger role in the overall carbon balance. The use in the LEA of observationally based estimates of heterotrophic respiration and fire disturbance, along with a more complete description of disturbance fluxes, suggests that the partitioning derived by the LEA is more likely, and that improved process descriptions and constraints by data are needed in the DGVMs.
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abstractUsing light aircraft and at intervals of approximately 14 days, vertical profiles of temperature, humidity, CO2 concentration and 13C/12C and 18O/16O ratio, as well as concentrations of CH4, CO, H2 and N2O, from about 80 to 3000 m above ground level have been determined for the atmosphere above a flux measurement tower located near the village of Zotino in central Siberia (60°45′N, 89°23′E). As well as being determined from flask measurements (typically at heights of 100, 500, 1000, 1500, 2000, 2500 and 3000 m) continuous CO2 concentration profiles at 1 Hz have also been obtained using an infrared gas analyser. This measurement program is ongoing and has been in existence since July 1998. Data to November 2000 are presented and show a seasonal cycle for CO2 concentration of about 25 μmol mol−1 within the atmospheric boundary layer (ABL) and about 15 μmol mol−1 in the free troposphere. Marked seasonal cycles in the isotopic compositions of CO2 are also observed, with that of oxygen-18 in CO2 being unusual: always being depleted in the ABL with respect to the free troposphere above. This is irrespective of whether the CO2 concentration is higher or lower in the free troposphere. We interpret this as indicating a net negative discrimination being associated with the net terrestrial carbon exchange, irrespective of whether photosynthesis or respiration dominates the net carbon flux in this region. During winter flights, large fluctuations in CO2 concentration with height are often observed both within and above the stable ABL. Usually (but not always) these variations in CO2 concentrations are associated with more or less stoichiometrically constant variations in CO and CH4 concentrations. We interpret this as reflecting the frequent transport of polluted air from Europe with very little vertical mixing having occurred, despite the large horizontal distances traversed. This notion is supported by back-trajectory analyses. Vertical profiles of CO2 concentration with supplementary flask measurements allow more information on the structure and composition of an air mass to be obtained than is the case for flask measurements or for ground-based measurements only. In particular, our data question the notion that there is usually anything like “well mixed background air” in the mid-to-high northern latitudes during the winter months.
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abstractWe present a first analysis of data (June 1998 to December 2000) from the long-term eddy covariance site established in a Pinus sylvestris stand near Zotino in central Siberia as part of the EUROSIBERIAN CARBONFLUX project. As well as examining seasonal patterns in net ecosystem exchange (NE), daily, seasonal and annual estimates of the canopy photosynthesis (or gross primary productivity, GP) were obtained using NE and ecosystem respiration measurements.Although the forest was a small (but significant) source of CO2 throughout the snow season (typically mid-October to early May) there was a rapid commencement of photosynthetic capacity shortly following the commencement of above-zero air temperatures in spring: in 1999 the forest went from a quiescent state to significant photosynthetic activity in only a few days. Nevertheless, canopy photosynthetic capacity was observed to continue to increase slowly throughout the summer months for both 1999 and 2000, reaching a maximum capacity in early August. During September there was a marked decline in canopy photosynthesis which was only partially attributable to less favourable environmental conditions. This suggests a reduction in canopy photosynthetic capacity in autumn, perhaps associated with the cold hardening process. For individual time periods the canopy photosynthetic rate was mostly dependent upon incoming photon irradiance. However, reductions in both canopy conductance and overall photosynthetic rate in response to high canopy-to-air vapour differences were clearly evident on hot dry days. The relationship between canopy conductance and photosynthesis was examined using Cowan's notion of optimality in which stomata serve to maximise the marginal evaporative cost of plant carbon gain. The associated Lagrangian multiplier (λ) was surprisingly constant throughout the growing season. Somewhat remarkably, however, its value was markedly different between years, being 416 mol mol−1 in 1999 but 815 mol mol−1 in 2000. Overall the forest was a substantial sink for CO2 in both 1999 and 2000: around 13 mol C m−2 a−1. Data from this experiment, when combined with estimates of net primary productivity from biomass sampling suggest that about 20% of this sink was associated with increasing plant biomass and about 80% with an increase in the litter and soil organic carbon pools. This high implied rate of carbon accumulation in the litter soil organic matter pool seems unsustainable in the long term and is hard to explain on the basis of current knowledge.
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Results of measurements and calculations of carbon budget parameters of forests and swamps of Siberia are reported. The zonal variability of reserves (and an increment in reserves) of carbon in forest and swamp ecosystems is characterized, carbon dioxide fluxes are measured directly by means of microeddy pulsations, and an uncertainty brought into the calculation of carbon budget parameters by forest fires is estimated.
Chapter
Active as well as passive spaceborne sensors can be used to monitor spring snowmelt on regional to continental scale. Change detection methods are used to determine dates related to the thaw period. They comprise initial thaw, primary thaw, start of diurnal thaw/refreeze period, mean date of thaw, end of thaw and start of greening-up of vegetation. Only the latter is determined by use of passive optical sensors and combines measurements of visible and infrared radiation. All other approaches use microwave data. Some instruments such as the scatterometer Seawinds on QuikScat and the radiometer AMSR-E on Aqua make several measurements per day allowing the detection of diurnal thaw and refreeze, which is characteristic of the spring snowmelt period in northern latitudes. A specific, diurnal difference approach developed for QuikScat allows the determination of the length of the final period of diurnal thaw/refreeze. This duration and the spatial dynamics are closely linked to surface hydrology and ecosystem processes. KeywordsSnowmelt-Active microwaves-Satellite data
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Carbon dioxide exchange between the ecosystem and the atmosphere would be a major component for carbon budget at boreal forests. In this chapter, net ecosystem exchange (NEE) of CO2 at a permafrost larch ecosystem will be discussed, based on a micrometeorological (tower flux) measurement. Movement of CO2 from ecosystem to atmosphere is customarily labeled as positive. The micrometeorological measurement can obtain NEE with a half-hourly time-resolution at an ecosystem scale. These temporal and spatial scales are advantages in carbon, water, and energy budget studies over ecological measurements.
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Zonal patterns of above-ground phytomass dynamics and carbon storage in above-ground vegetation, phytodetritus and humus were revealed based on the study of the carbon balance and its components in forest ecosystems of the Yenisei meridian transect. Results indicate that the carbon storage ratio in different plant layers is determined by climatic regimes. For example pine stands were used to calculate the full carbon budget using data on its fluxes and storage in different biogeocenosis components. Biological productivity indices and the carbon pool of hydromorphic complexes are highly dependant on the mineral nutrition regime and morphological characteristics of the stands. Experimental study results show the importance of forest and bog ecosystems as carbon cycle regulators is determined by the complex interaction of zonal-climatic and forest conditions as well as by forest vegetation characteristics (which depend on varying carbon balance structure and energy- mass exchange processes).
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We have built the first comprehensive global three-dimensional model of δ18O in atmospheric CO2. The constructed model goes beyond all other approaches made until now, by simulating the diurnal variations and transport of CO2, δ18O of water, and δ18O of CO2. The CO18O fluxes are thereby dependent on the atmospheric CO18O composition. We have validated the model surface processes, showing that it compares well to other estimates and measurements of NPP, NEE, and stomata-internal CO2 mixing ratio (ci), except for high northern latitudes. Here, the model is considerably lower in NPP and higher in ci than other model estimates. However, estimates derived indirectly from observations tend to support our model findings. The water isotopes of rain are reproduced very well at all latitudes. The soil bucket model used in the model integrates incoming rain in one single value. The bucket approach overattenuates the isotopic variations of rain, and hence our isotopic source signature of respiration shows almost no seasonal cycle and is thus isotopically too depleted during summer.
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The spatial upscaling of soil respiration from field measurements to ecosystem levels will be biased without studying its spatial variation. We took advantage of the unique spatial gradients of an oak–grass savanna ecosystem in California, with widely spaced oak trees overlying a grass layer, to study the spatial variation in soil respiration and to use these natural gradients to partition soil respiration according to its autotrophic and heterotrophic components. We measured soil respiration along a 42.5 m transect between two oak trees in 2001 and 2002, and found that soil respiration under tree canopies decreased with distance from its base. In the open area, tree roots have no influence on soil respiration. Seasonally, soil respiration increased in spring until late April, and decreased in summer following the decrease in soil moisture content, despite the further increase in soil temperature. Soil respiration significantly increased following the rain events in autumn. During the grass growing season between November and mid-May, the average of CO2 efflux under trees was 2.29 µmol m-2 s-1, while CO2 efflux from the open area was 1.40 µmol m-2 s-1. We deduced that oak root respiration averaged as 0.89 µmol m-2 s-1, accounting for 39% of total soil respiration (oak root + grass root + microbes). During the dry season between mid-May and October, the average of CO2 efflux under trees was 0.87 µmol m-2 s-1, while CO2 efflux from the open areas was 0.51 µmol m-2 s-1. Oak root respiration was 0.36 µmol m-2 s-1, accounting for 41% of total soil respiration (oak root + microbes). The seasonal pattern of soil CO2 efflux under trees and in open areas was simulated by a bi-variable model driven by soil temperature and moisture. The diurnal pattern was influenced by tree physiology as well. Based on the spatial gradient of soil respiration, spatial analysis of crown closure and the simulation model, we spatially and temporally upscaled chamber measurements to the ecosystem scale. We estimated that the cumulative soil respiration in 2002 was 394 gC m-2 year-1 in the open area and 616 gC m-2 year-1 under trees with a site-average of 488 gC m-2 year-1.
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INTRODUCTION Soil respiration is a major component in the carbon balance of terrestrial ecosystems and has been measured in the field for more than eight decades. In this chapter, we will describe the measurement of soil CO2 efflux at the soil surface that can be considered as equivalent to soil CO2 production when integrated over long time periods (week, month or season). At shorter time scales the transport of CO2 may uncouple the soil CO2 efflux from its production inside the soil. Different methods have been developed to measure this efflux. These methods can affect the object being measured by disturbing the biochemical processes involved in CO2 production, the physical properties influencing CO2 movement towards the soil surface, or by changing the environmental conditions in the soil. Therefore, soil respiration measurements in the field are one of the most difficult among the ecosystem flux measurements. So far, no single method has been established as the standard but comparisons, which give important indications on their accuracy, have been performed. The choice of the measurement methodology is not limited to that of a measurement system. The experimenter has to elaborate a protocol depending on the temporal and spatial scales studied. In this chapter, we will describe the most commonly used methodologies for measuring soil CO2 efflux and present their history, principles and constraints (Section 2.2).
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A range of observations points towards earlier onset of spring in northern high latitudes. However, despite the profound effects this may have on vegetation-atmosphere exchange of carbon (NEE), vegetation-atmosphere physical coupling, or the location of the tundrataiga interface, the number of studies that investigate winter-spring transition fluxes in contrasting northern vegetation types is limited. Here, we examine spring ecosystem-atmosphere energy and carbon exchange in a Siberian pine forest and mire. Divergent surface albedo before and during snow-melt resulted in daytime net radiation (R n) above the forest exceeding Rn above the mire by up to 10 MJ m-2. Until stomata could open, absorbed radiation by the green pine canopy caused substantial daytime sensible heat fluxes (H > 10MJ m -2). H above the mire was very low, even negative (< -2 MJ m -2), during that same period. Physiological activity in both ecosystems responded rapidly to warming temperatures and snow-melt, which is essential for survival in Siberia with its very short summers. On days with above-zero temperatures, before melt was complete, low rates of forest photosynthesis (1-2 μmol m-2 s-1) were discernible. Forest and mire NEE became negative the same day, or shortly after, photosynthesis commenced. The mire lagged by about two weeks behind the forest and regained its full carbon uptake capacity at a slower rate. Our data provide empirical evidence for the importance the timing of spring and the relative proportion of forest vs. mire has for late winter/spring boundary-layer growth, and production and surface-atmosphere mixing of trace gases. Models that seek to investigate effects of increasingly earlier spring in high latitudes must correctly account for contrasting physical and biogeochemical ecosystem-atmosphere exchange in heterogeneous landscapes.
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We compare assimilation and respiration rates, and water use strategies in four divergent ecosystems located in cold-continental central Siberia and in semi-arid southern Africa. These seemingly unrelated systems have in common a harsh and highly seasonal environment with a very sharp transition between the dormant and the active season, with vegetation facing dry air and soil conditions for at least part of the year. Moreover, the northern high latitudes and the semi-arid tropics will likely experience changes in key environmental parameters (e.g., air temperature and precipitation) in the future; indeed, in some regions marked climate trends have already been observed over the last decade or so.
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Stem maintenance respiration was linearly related to live-cell volume for lodgepole pine from 4-36 cm dbh and for Engelmann spruce from 0-20 cm dbh. Sapwood contained >80% of the total live-cell volume in stems. Bole surface area, commonly used to estimate tree respiration costs, poorly estimated stem maintenance respiration. At 15°C, maintenance costs for lodgepole pine were 6.6 × 10-5 kg C.(Kg C sapwood)-1.d-1. Stem respiration during the growing season correlated well with annual stemwood growth. Annual stem maintenance respiration for trees and stand can be estimated using sapwood volume, sapwood temperature, and knowledge of respiratory behaviour. Total respiration (construction plus maintenance) estimated using stem growth and a model of maintenance respiration was compared with actual respiration measurements integrated over a 100-d growing season. Estimated respiration agreed with the integrated measurements for Engelmann spruce, but overestimated the integrated measurements by 73% in lodgepole pine. -from Authors
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Past studies of plant-microbe interactions in the alpine nitrogen cycle have revealed a seasonal separation of N use, with plants absorbing N primarily during the summer months and microbes immobilizing N primarily during the autumn months. On the basis of these studies, it has been concluded that competition for N between plants and microbes is minimized along this seasonal gradient. In this study, we examined more deeply the links between microbial population dynamics and plant N availability in an alpine dry meadow. We conducted a year-round field study and performed experiments on isolated soil microorganisms. Based on previous work in this ecosystem, we hypothesized that microbial biomass would decline before the plant growing season and would release N that would become available to plants. Microbial biomass was highest when soils were cold, in autumn, winter, and early spring. During this time, N was immobilized in microbial biomass. After snow melt in spring, microbial biomass decreased. A peak in the soil protein concentration was seen at this time, followed by peaks in soil amino acid and ammonium concentrations in late June. Soil protease rates were initially high after snow melt, decreased to below detection limits by midsummer, and partially recovered by late summer. Proteolytic activity in soil was saturated early in the growing season and became protein limited later in the summer. We concluded that the key event controlling N availability to alpine plants occurs after snow melt, when protein is released from the winter microbial biomass. This protein pulse provides substrate for soil proteases, which supply plants with amino acids during the growing season. On average, microbial biomass was lower in the summer than at other times, although the biomass fluctuated widely during the summer. Within the summer months, maximum numbers of amino-acid-degrading microorganisms and the maximum amount of microbial biomass coincided with the peak in soil amino acids, when plants are most active. All bacterial strains isolated from this summer community had the ability to grow rapidly on low concentrations of amino acids and to degrade protein. This explains the previously observed result that the soil microbial biomass can compete strongly with plants for organic N, despite the seasonal offset of maximum plant and microbial N uptake.
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Autotrophic respiration (Ra) in forest ecosystems can be >50% of the carbon fixed in photosynthesis and may regulate productivity and carbon storage in forest ecosystems, because Ra increases with temperature. We estimated annual Ra from chamber measurements in aspen, black spruce, and jack pine forests in Canada for 1994. Mean foliage respiration at 10°C for expanded leaves was 0.21–0.95 μmol m−2 (leaf surface) s−1 for all species and differed little from May to September. Wood respiration at 15°C (0.2–1 μmol m−2 (stem surface) s−1 for all species) was strongly seasonal, with high rates in midsummer that coincided with wood growth. Fine root respiration at 10°C was 2.5–7.7 μmol kg−1 s−1 for all species and declined throughout the growing season for the conifers. Annual costs of Ra for foliage, wood, and roots (overstory and understory) were 490, 610, and 450 g C m−2 (ground) yr−1 for aspen, black spruce, and jack pine (old) in northern Manitoba and 600, 480, and 310 g C m−2 yr−1 for aspen, black spruce, and jack pine (old) in central Saskatchewan. Carbon use efficiency (CUE), the ratio of net production to production plus Ra, averaged 0.44, 0.34, and 0.39 for aspen, black spruce, and jack pine (old) for all tissues and 0.61, 0.36, and 0.44 for aboveground tissues. Differences in CUE between the northern and the southern sites were small for all species, and CUE did not vary with stand biomass. Species differences in CUE suggest that models assuming a constant CUE across species may poorly estimate production and carbon balance for any given site.
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We investigated the effect of temperature and irradiance on leaf respiration (R, non-photorespiratory mitochondrial CO(2) release) of snow gum (Eucalyptus pauciflora Sieb. ex Spreng). Seedlings were hydroponically grown under constant 20 degrees C, controlled-environment conditions. Measurements of R (using the Laisk method) and photosynthesis (at 37 Pa CO(2)) were made at several irradiances (0-2,000 micromol photons m(-2) s(-1)) and temperatures (6 degrees C-30 degrees C). At 15 degrees C to 30 degrees C, substantial inhibition of R occurred at 12 micromol photons m(-2) s(-1), with maximum inhibition occurring at 100 to 200 micromol photons m(-2) s(-1). Higher irradiance had little additional effect on R at these moderate temperatures. The irradiance necessary to maximally inhibit R at 6 degrees C to 10 degrees C was lower than that at 15 degrees C to 30 degrees C. Moreover, although R was inhibited by low irradiance at 6 degrees C to 10 degrees C, it recovered with progressive increases in irradiance. The temperature sensitivity of R was greater in darkness than under bright light. At 30 degrees C and high irradiance, light-inhibited rates of R represented 2% of gross CO(2) uptake (v(c)), whereas photorespiratory CO(2) release was approximately 20% of v(c). If light had not inhibited leaf respiration at 30 degrees C and high irradiance, R would have represented 11% of v(c). Variations in light inhibition of R can therefore have a substantial impact on the proportion of photosynthesis that is respired. We conclude that the rate of R in the light is highly variable, being dependent on irradiance and temperature.
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The turbulent exchanges of CO2 and water vapour between an aggrading deciduous forest in the north-eastern United States (Harvard Forest) and the atmosphere were measured from 1990 to 1994 using the eddy covariance technique. We present a detailed description of the methods used and a rigorous evaluation of the precision and accuracy of these measurements. We partition the sources of error into three categories: (1) uniform systematic errors are constant and independent of measurement conditions (2) selective systematic errors result when the accuracy of the exchange measurement varies as a function of the physical environment, and (3) sampling uncertainty results when summing an incomplete data set to calculate long-term exchange. Analysis of the surface energy budget indicates a uniform systematic error in the turbulent exchange measurements of -20 to 0%. A comparison of nocturnal eddy flux with chamber measurements indicates a selective systematic underestimation during calm (friction velocity < 0.17 m s−1) nocturnal periods. We describe an approach to correct for this error. The integrated carbon sequestration in 1994 was 2.1 t C ha−1 y−1 with a 90% confidence interval due to sampling uncertainty of ±0.3 t C ha−1 y−1 determined by Monte Carlo simulation. Sampling uncertainty may be reduced by estimating the flux as a function of the physical environment during periods when direct observations are unavailable, and by minimizing the length of intervals without flux data. These analyses lead us to place an overall uncertainty on the annual carbon sequestration in 1994 of -0.3 to +0.8 t C ha−1 y−1.
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Soil respiration includes soil microbial respiration, soil fauna respiration, and plant root respiration, therefore it reflects the biological activity of the soil ecosystems. The Siberian Taiga often experiences serious damage from forest fire, due to the very low precipitation in spring. We measured the soil respiration in five forest soil ecosystems with different histories of forest fire in Yakutsk in August 1997. The dominant tree species was Larix cajanderi, and the soils were Spodosols with a sandy and loamy texture. We also measured the soil respiration in a grassland. At severely burned sites, almost all the trees had fallen, litter and vegetation on the forest floor had burned, other forms of vegetation, including bryophytes or herbs, had invaded. At less severely burned forest sites, the trees were still standing but litter and vegetation on the forest floor had disappeared. Soil temperature, moisture, pH, and EC all increased after severe forest fires, the A-horizon showed a higher organic carbon content and a lower CN ratio. Soil respiration rate ranged from 18 to 397 (10 g CO2 m ) in the same order reported so far. Soil respiration in severely burned forests was significantly lower than in intact forests, and was similar to that of grassland. Furthermore, mildly burned forests showed soil respiration values intermediate between those of severely burned and intact forests. These findings suggest that tree root respiration is considerably higher than root respiration of other plants or microbial and fauna respiration in soil. Soil microbial respiration was determined by the incubation method under the same temperature and soil moisture conditions as those in situ. Multiple regression analysis for mineral soils showed that the soil microbial respiration increased with the increase of the soil temperature and organic carbon content, that the soil microbial respiration decreased with the increase of pH. Whole soil microbial respiration within 1 m depth was higher in severely burned forests than in intact forests. These findings show that forest fire increased the soil microbial respiration and confirm that the loss of tree root respiration was the main reason for the decrease in soil respiration after severe forest fire. The contribution of tree root respiration to soil respiration was estimated to exceed 50%. Severe forest fire kills trees, and consequently results in a decrease of soil respiration.
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The causes of a reduced sensitivity of high-latitude tree growth to variations in summer temperature for recent decades,, compared to earlier this century, are unknown. This sensitivity change is problematic, in that relationships between tree-ring properties and temperature are widely used for reconstructing past climate. Here we report an analysis of tree-ring and climate data from the forest-tundra zone, in combination with a mechanistic model of tree-ring growth, to argue that an increasing trend of winter precipitation over the past century in many subarctic regions led to delayed snow melt in these permafrost environments. As a result, the initiation of cambial activity (necessary for the formation of wood cells) has been delayed relative to the pre-1960 period in the Siberian subarctic. Since the early 1960s, less of the growth season has been during what had previously been the period of maximal growth sensitivity to temperature. This shift results not only in slower growth, but also in a reduced correlation between growth and temperature. Our results suggest that changes in winter precipitation should be considered in seeking explanations for observed changes in the timing of the `spring greening' of high-latitude forests, and should be taken into account in the study of the role of the Siberian subarctic forest in the global carbon cycle.
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The objectives of this study are to (1) characterize the carbon (C) content, leaf area index, and aboveground net primary production (ANPP) for mature aspen, black spruce, and young and mature jack pine stands at the southern and northern Boreal Ecosystem-Atmosphere Study (BOREAS) areas and (2) compare net primary production and carbon allocation coefficients for the major boreal forest types of the world. Direct estimates of leaf area index, defined as one half of the total leaf surface area, range from a minimum of 1.8 for jack pine forests to a maximum of 5.6 for black spruce forests; stems comprise 5 to 15% of the total overstory plant area. In the BOREAS study, total ecosystem (vegetation plus detritus plus soil) carbon content is greatest in the black spruce forests (445,760-479,380kgCha-1), with 87 to 88% of the C in the soil, and is lowest in the jack pine stands (68,370-68,980kgCha-1) with a similar distribution of carbon in the vegetation and soil. Forest floor carbon content and mean residence time (MRT) also vary more among forest types in a study area than between study areas for a forest type; forest floor MRT range from 16 to 19 years for aspen stands to 28 to 39 years for jack pine stands. ANPP differs significantly among the mature forests at each of the BOREAS study areas, ranging from a maximum of 3490 to 3520kgCha-1yr-1 for aspen stands to 1170 to 1220kgCha-1yr-1 for jack pine stands. Both net primary production (NPP) and carbon allocation differ between boreal evergreen and deciduous forests in the world, suggesting global primary production models should distinguish between these two forest types. On average, 56% of NPP for boreal forests occurs as detritus and illustrates the need to better understand factors controlling aboveground and below-ground detritus production in boreal forests.
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We use semi-mechanistic, empirically based statistical models to predict the spatial and temporal patterns of global carbon dioxide emissions from terrestrial soils. Emissions include the respiration of both soil organisms and plant roots. At the global scale, rates of soil CO2 efflux correlate significantly with temperature and precipitation; they do not correlate well with soil carbon pools, soil nitrogen pools, or soil C:N. Wetlands cover about 3% of the land area but diminish predicted CO2 emissions by only about 1%. The estimated annual flux of CO2 from soils to the atmosphere is estimated to be 76.5 Pg C yr−1, 1–9 Pg greater than previous global estimates, and 30–60% greater than terrestrial net primary productivity. Historic land cover changes are estimated to have reduced current annual soil CO2 emissions by 0.2–2.0 Pg C yr−1 in comparison with an undisturbed vegetation cover. Soil CO2 fluxes have a pronounced seasonal pattern in most locations, with maximum emissions coinciding with periods of active plant growth. Our models suggest that soils produce CO2 throughout the year and thereby contribute to the observed wintertime increases in atmospheric CO2 concentrations. Our derivation of statistically based estimates of soil CO2 emissions at a 0.5° latitude by longitude spatial and monthly temporal resolution represents the best-resolved estimate to date of global CO2 fluxes from soils and should facilitate investigations of net carbon exchanges between the atmosphere and terrestrial biosphere.
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Predicted daily fluxes from an ecosystem model for water, carbon dioxide, and methane were compared with 1994 and 1996 Boreal Ecosystem-Atmosphere Study (BOREAS) field measurements at sites dominated by old black spruce (Picea mariana (Mill.) BSP) (OBS) and boreal fen vegetation near Thompson, Man. Model settings for simulat- ing daily changes in water table depth (WTD) for both sites were designed to match observed water levels, including predictions for two microtopographic positions (hollow and hummock) within the fen study area. Water run-on to the soil profile from neighboring microtopographic units was calibrated on the basis of daily snowmelt and rainfall inputs to reproduce BOREAS site measurements for timing and magnitude of maximum daily WTD for the growing season. Model predictions for daily evapotranspiration rates closely track measured fluxes for stand water loss in patterns con- sistent with strong controls over latent heat fluxes by soil temperature during nongrowing season months and by vari- ability in relative humidity and air temperature during the growing season. Predicted annual net primary production (NPP) for the OBS site was 158 g C·m-2 during 1994 and 135 g C·m-2 during 1996, with contributions of 75% from overstory canopy production and 25% from ground cover production. Annual NPP for the wetter fen site was 250 g C·m-2 during 1994 and 270 g C·m-2 during 1996. Predicted seasonal patterns for soil CO 2 fluxes and net ecosystem production of carbon both match daily average estimates at the two sites. Model results for methane flux, which also closely match average measured flux levels of -0.5 mg CH4·m-2·day-1 for OBS and 2.8 mg CH4·m-2·day-1 for fen sites, suggest that spruce areas are net annual sinks of about -0.12 g CH4·m-2, whereas fen areas generate net annual emis- sions on the order of 0.3-0.85 g CH4·m-2, depending mainly on seasonal WTD and microtopographic position. Fen hollow areas are predicted to emit almost three times more methane during a given year than fen hummock areas. The validated model is structured for extrapolation to regional simulations of interannual trace gas fluxes over the entire North America boreal forest, with integration of satellite data to characterize properties of the land surface.
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TURC, a diagnostic model for the estimation of continental gross primary productivity (GPP) and net primary productivity (NPP), is presented. This model uses a remotely sensed vegetation index to estimate the fraction of solar radiation absorbed by canopies, and an original parameterization of the relationship between absorbed solar radiation and GPP, based on measurements of CO2 fluxes above plant canopies. An independent, uncalibrated model of autotrophic maintenance and growth respiration is parameterized from literature data, and uses databases on temperature, biomass, and remotely sensed vegetation index. This model results in global estimates of GPP and NPP of 133.1 and 62.3 Gt(C) per year, respectively, which is consistent with commonly admitted values. The ratio of autotrophic respiration to GPP is about 70% for equatorial rain forests and 50% for temperate forests, as a result the highest predicted NPP are in tropical savannas of Africa and South America, and in temperate, highly cultivated zones of North America, not in equatorial rain forest zones. Conversion efficiencies defined as the ratio of yearly integrated NPP to absorbed photosynthetically active radiation (PAR) compare relatively well with a previous compilation of literature values, except for ecosystems with probable reduction of conversion efficiency due to water stress. Several sensitivity studies are performed on some input data sets, model assumptions, and model parameters.
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The term zero curtain refers to the effect of latent heat in maintaining temperatures near 0°C over extended periods in freezing or thawing soils. Analysis of thermal data from an arctic site and a mid-latitude location suggests that the phenomenon may be a universal characteristic of wet, medium-textured mineral and organic soils during freezing. Nonconductive processes play an important role in formation and maintenance of the zero-curtain effect. Water advection, condensation, and evaporation events are recognizable in mappings of thermal and ion concentration records on time-depth meshes. The zero curtain appears to be produced and maintained by vapor transport and internal distillation mechanisms, driven by osmotic potential variations induced by freeze-thaw events at the soil surface. The term zero curtain is applicable to locations outside permafrost regions, but units of measurements should not be assigned to it.
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Abstract We present results from two years’ net ecosystem flux measurements above a boreal forest in central Sweden. Fluxes were measured with an eddy correlation system based on a sonic anemometer and a closed path CO2 and H2O gas analyser. The measurements show that the forest acted as a source during this period, and that the annual balance is highly sensitive to changes in temperature. The accumulated flux of carbon dioxide during the full two-year period was in the range 480–1600 g CO2 m–2. The broad range is caused by uncertainty regarding assessment of the night-time fluxes. Although annual mean temperature remained close to normal, the results are partly explained by higher than normal respiration, due to abnormal temperature distribution and reduced soil moisture during one growing season. The finding that a closed forest can be a source of carbon over such a long period as two years contrasts sharply with the common belief that forests are always carbon sinks.
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Modeling the terrestrial biosphere's carbon exchanges constitutes a key tool for investigation of the global carbon cycle, which has lead to the recent development of numerous terrestrial biosphere models. However, as demonstrated by recent intercomparison studies, results of plant carbon uptake, expressed as net primary productivity (NPP), still diverge to a large degree. Here, we address the question of uncertainty by conducting a series of sensitivity tests with a single, process-based model, the Biosphere Energy-Transfer Hydrology (BETHY) scheme. We calculate NPP globally for a standard model setup and various alternative model setups representing either changes in modeling strategy or approximate uncertainties of the most important model parameters. The results show that estimated uncertainties of many process parameters are still too large for reliable predictions of global NPP. The largest uncertainties come from plant respiration, photosynthesis and soil water storage. The surface radiation balance and day-to-day variations in weather, often not included into terrestrial vegetation models, are also found to contribute significantly to overall uncertainties, while stomatal behavior, the aerodynamic coupling of vegetation and atmosphere, and the choice of the vegetation map turn out to be relatively unimportant. A further comparison with field measurements of NPP suggests that such data are too unreliable for validating biosphere modél predictions. We conclude that the inherent uncertainties in process-oriented biosphere modeling are able to explain the discrepancies that have occurred when comparing the results of different models.
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Initial observations of the diurnal variation of the vertical thermal structure of a loblolly pine planation are presented. Results obtained in other forests are qualitatively confirmed. On a clear day a very unstable temperature gradient occurs above the trees, while a strong inversion (∼8°C) develops below the crowns. At night the sub-crown region becomes weakly unstable, but the atmospheric layer above the trees is then stable. On a rainy day, the strength of the temperature inversion beneath the tree crowns is less than 1°C. The position of the daytime temperature maximum in the tree-tops responds to the solar elevation, eventually descending about 2 m to the region of maximum foliage density as the sun's rays penetrate deeper into the tree crowns.
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Estimates of the extent of the discrimination against13CO2 during photosynthesis (?A) on a global basis were made using gridded data sets of temperature, precipitation, elevation, humidity and vegetation type. Stomatal responses to leaf-to-air vapour mole fraction difference (D, leaf-to-air vapour pressure difference divided by atmospheric pressure) were first determined by a literature review and by assuming that stomatal behaviour results in the optimisation of plant water use in relation to carbon gain. Using monthly time steps, modelled stomatal responses toD were used to calculate the ratio of stomatal cavity to ambient CO2 mole fractions and then, in association with leaf internal conductances, to calculate ?A. Weighted according to gross primary productivity (GPP, annual net CO2 asimilation per unit ground area), estimated ?A for C3 biomes ranged from 12.9‰ for xerophytic woods and shrub to 19.6‰ for cool/cold deciduous forest, with an average value from C3 plants of 17.8‰. This is slightly less than the commonly used values of 18–20‰. For C4 plants the average modelled discrimination was 3.6‰, again slightly less than would be calculated from C4 plant dry matter carbon isotopic composition (yielding around 5‰). From our model we estimate that, on a global basis, 21% of GPP is by C4 plants and for the terrestrial biosphere as a whole we calculate an average isotope discrimination during photosynthesis of 14.8‰. There are large variations in ?A across the globe, the largest of which are associated with the precence or absence of C4 plants. Due to longitudinal variations in ?A, there are problems in using latitudinally averaged terrestrial carbon isotope discriminations to calculate the ratio of net oceanic to net terrestrial carbon fluxes.
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The exchange of carbon dioxide (CO2) between the atmosphere and terrestrial ecosystems due to photosynthesis and respiration has been simulated using a new version of the simple biosphere model (SiB2) and the Colorado State University (CSU) general circulation model (GCM). Parameters associated with the extent and seasonality of vegetation were derived from satellite observations. The fluxes were calculated at the GCM time step of 6 min, so that the diurnal cycle of photosynthesis is well resolved. Annual net primary productivity simulated by the coupled model agrees well with previous estimates in most regions of the world. In some regions (central North America, southeastern South America, southeast Asia), the precipitation simulated by the CSU GCM is less than observed, and in those regions the simulated NPP is less than previous estimates. The amplitude of the seasonal cycle of the simulated net flux is quite similar to previous estimates, but the phase is significantly earlier in the northern temperate and boreal zones, both as simulated by the GCM and when SiB2 is driven off-line using observed meteorological forcing. At the few locations for which observational data are available, the phase of the simulated seasonal cycle of net carbon fluxes agrees well with the data, but at one temperate forest grid cell the amplitude is too low. The phase of the simulated diurnal cycle reproduces observations from a temperate forest, temperate grassland, and tropical forest. The amplitude of the simulated diurnal cycle is close to the observed amplitude early in the season at the temperate grassland site, but deteriorates by late summer because of drought stress due to the less-than-observed simulated precipitation.
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Abstract Quantum yields of photosynthetic CO2 uptake by Pinus sylvestris (L.) shoots were measured at temperatures between −; 2°C and 35°C from September 1984 to September 1985. The ratio of variable to peak fluorescence of photosystem II (Fv/Fp) was also measured. Quantum yield measured at 25°C varied with time from a low winter value of 0.017 to a high summer value of 0.057. This variation was strongly correlated to variation in FvFp(r2= 0.91). The response of quantum yield to temperature changed with season. During winter, quantum yield was essentially constant between 0°C and 35°C. The constancy above 5°C was associated with a strong increase in intercellular space CO2 (Ci) with temperature. In June, the quantum yield peaked at 5°C, decreased sharply below 5°C, and was rather constant between 25°C and 35°C. This insensitivity to increased temperature above 25°C was attributed to a large increase in Ci In contrast, by September, the quantum yield was less sensitive to temperature below 5°C and more sensitive above 25°C, despite an unchanged Ci response with increasing temperatures as compared with June. In August, quantum yields were lowered at 0, 5 and 15°C, apparently as a result of high carbohydrate levels in the leaves. Overall, the results suggest that there are sites other than in photosystem II or at ribulose bisphosphate carboxylase/oxygenase at which the quantum yield of photosynthetic CO2 uptake is affected. Possible causes for the changes in efficiency are discussed.
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The effects of fertilization, irrigation or both on the seasonal changes of starch and soluble carbohydrates (glucose, fructose, myo-inositol, pinitol and sucrose) in needles of 20-year-old Scots pine trees (Pinus silvestris L.) were studied during three consecutive years. The starch content of the mature needles increased during spring and early summer to about 25% of dry weight. Neither fertilization nor irrigation affected the general pattern of starch accumulation during the spring. The starch reserves were mobilized when the shoot started to grow. Starch content decreased more rapidly in needles from fertilized than in those from unfertilized trees. The current needles from the control trees accumulated starch while they were still growing. The current needles of the fertilized trees did so to a lesser extent. The amount of starch was closely correlated to the air temperature and to the growth rate. Large amounts were found at low temperatures and low growth rates. The concentrations of soluble carbohydrates showed the well-known seasonal variation, with the highest value during the winter. The levels of sugars were nearly similar, irrespective of fertilization. An exception was sucrose, which was found in small quantities in needles from fertilized plots. Small amounts of sucrose were also found in growing current needles. The results are discussed in relation to growth limitation by assimilate availability and indicate that the ‘sink demand’ is the limiting factor.
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The possible responses of ecosystem processes to rising atmospheric CO2 concentration and climate change are illustrated using six dynamic global vegetation models that explicitly represent the interactions of ecosystem carbon and water exchanges with vegetation dynamics. The models are driven by the IPCC IS92a scenario of rising CO2 (Wigley et al. 1991), and by climate changes resulting from effective CO2 concentrations corresponding to IS92a, simulated by the coupled ocean atmosphere model HadCM2-SUL. Simulations with changing CO2 alone show a widely distributed terrestrial carbon sink of 1.4–3.8 Pg C y−1 during the 1990s, rising to 3.7–8.6 Pg C y−1 a century later. Simulations including climate change show a reduced sink both today (0.6–3.0 Pg C y−1) and a century later (0.3–6.6 Pg C y−1) as a result of the impacts of climate change on NEP of tropical and southern hemisphere ecosystems. In all models, the rate of increase of NEP begins to level off around 2030 as a consequence of the ‘diminishing return’ of physiological CO2 effects at high CO2 concentrations. Four out of the six models show a further, climate-induced decline in NEP resulting from increased heterotrophic respiration and declining tropical NPP after 2050. Changes in vegetation structure influence the magnitude and spatial pattern of the carbon sink and, in combination with changing climate, also freshwater availability (runoff). It is shown that these changes, once set in motion, would continue to evolve for at least a century even if atmospheric CO2 concentration and climate could be instantaneously stabilized. The results should be considered illustrative in the sense that the choice of CO2 concentration scenario was arbitrary and only one climate model scenario was used. However, the results serve to indicate a range of possible biospheric responses to CO2 and climate change. They reveal major uncertainties about the response of NEP to climate change resulting, primarily, from differences in the way that modelled global NPP responds to a changing climate. The simulations illustrate, however, that the magnitude of possible biospheric influences on the carbon balance requires that this factor is taken into account for future scenarios of atmospheric CO2 and climate change.
Article
Based on review and original data, this synthesis investigates carbon pools and fluxes of Siberian and European forests (600 and 300 million ha, respectively). We examine the productivity of ecosystems, expressed as positive rate when the amount of carbon in the ecosystem increases, while (following micrometeorological convention) downward fluxes from the atmosphere to the vegetation (NEE = Net Ecosystem Exchange) are expressed as negative numbers. Productivity parameters are Net Primary Productivity (NPP=whole plant growth), Net Ecosystem Productivity (NEP = CO 2 assimilation minus ecosystem respiration), and Net Biome Productivity (NBP = NEP minus carbon losses through disturbances bypassing respiration, e.g. by fire and logging). Based on chronosequence studies and national forestry statistics we estimate a low average NPP for boreal forests in Siberia: 123 gC m –2 y –1 . This contrasts with a similar calculation for Europe which suggests a much higher average NPP of 460 gC m –2 y –1 for the forests there. Despite a smaller area, European forests have a higher total NPP than Siberia (1.2–1.6 vs. 0.6–0.9 × 10 ¹⁵ gC region –1 y –1 ). This arises as a consequence of differences in growing season length, climate and nutrition. For a chronosequence of Pinus sylvestris stands studied in central Siberia during summer, NEE was most negative in a 67‐y old stand regenerating after fire (– 192 mmol m –2 d –1 ) which is close to NEE in a cultivated forest of Germany (– 210 mmol m –2 d –1 ). Considerable net ecosystem CO 2 ‐uptake was also measured in Siberia in 200‐ and 215‐y old stands (NEE:174 and – 63 mmol m –2 d –1 ) while NEP of 7‐ and 13‐y old logging areas were close to the ecosystem compensation point. Two Siberian bogs and a bog in European Russia were also significant carbon sinks (– 102 to – 104 mmol m –2 d –1 ). Integrated over a growing season (June to September) we measured a total growing season NEE of – 14 mol m –2 summer –1 (– 168 gC m –2 summer –1 ) in a 200‐y Siberian pine stand and – 5 mol m –2 summer –1 (– 60 gC m –2 summer –1 ) in Siberian and European Russian bogs. By contrast, over the same period, a spruce forest in European Russia was a carbon source to the atmosphere of (NEE: + 7 mol m –2 summer –1 = + 84 gC m –2 summer –1 ). Two years after a windthrow in European Russia, with all trees being uplifted and few successional species, lost 16 mol C m –2 to the atmosphere over a 3‐month in summer, compared to the cumulative NEE over a growing season in a German forest of – 15.5 mol m –2 summer –1 (– 186 gC m –2 summer –1 ; European flux network annual averaged – 205 gC m –2 y –1 ). Differences in CO 2 ‐exchange rates coincided with differences in the Bowen ratio, with logging areas partitioning most incoming radiation into sensible heat whereas bogs partitioned most into evaporation (latent heat). Effects of these different surface energy exchanges on local climate (convective storms and fires) and comparisons with the Canadian BOREAS experiment are discussed. Following a classification of disturbances and their effects on ecosystem carbon balances, fire and logging are discussed as the main processes causing carbon losses that bypass heterotrophic respiration in Siberia. Following two approaches, NBP was estimated to be only about 13–16 mmol m –2 y –1 for Siberia. It may reach 67 mmol m –2 y –1 in North America, and about 140–400 mmol m –2 y –1 in Scandinavia. We conclude that fire speeds up the carbon cycle, but that it results also in long‐term carbon sequestration by charcoal formation. For at least 14 years after logging, regrowth forests remain net sources of CO 2 to the atmosphere. This has important implications regarding the effects of Siberian forest management on atmospheric concentrations. For many years after logging has taken place, regrowth forests remain weaker sinks for atmospheric CO 2 than are nearby old‐growth forests.
Article
This paper presents CO2 flux data from 18 forest ecosystems, studied in the European Union funded EUROFLUX project. Overall, mean annual gross primary productivity (GPP, the total amount of carbon (C) fixed during photosynthesis) of these forests was 1380 ± 330 gC m−2 y−1 (mean ±SD). On average, 80% of GPP was respired by autotrophs and heterotrophs and released back into the atmosphere (total ecosystem respiration, TER = 1100 ± 260 gC m−2 y−1). Mean annual soil respiration (SR) was 760 ± 340 gC m−2 y−1 (55% of GPP and 69% of TER). Among the investigated forests, large differences were observed in annual SR and TER that were not correlated with mean annual temperature. However, a significant correlation was observed between annual SR and TER and GPP among the relatively undisturbed forests. On the assumption that (i) root respiration is constrained by the allocation of photosynthates to the roots, which is coupled to productivity, and that (ii) the largest fraction of heterotrophic soil respiration originates from decomposition of young organic matter (leaves, fine roots), whose availability also depends on primary productivity, it is hypothesized that differences in SR among forests are likely to depend more on productivity than on temperature. At sites where soil disturbance has occurred (e.g. ploughing, drainage), soil espiration was a larger component of the ecosystem C budget and deviated from the relationship between annual SR (and TER) and GPP observed among the less-disturbed forests. At one particular forest, carbon losses from the soil were so large, that in some years the site became a net source of carbon to the atmosphere. Excluding the disturbed sites from the present analysis reduced mean SR to 660 ± 290 gC m−2 y−1, representing 49% of GPP and 63% of TER in the relatively undisturbed forest ecosystems.
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The range and source of variation in foliage respiration rate in the dormant season were investigated for plants of Lycopodium annotinum L., Pinus contorta Dougl. var. latifolia Engelm., Picea abies (L.) Karst., Andromeda polifolia L., Calluna vulgaris (L.) Hull, Vaccinium myrtillus L., Vaccinium vitis-ideae L. and Empetrum hermaphroditum Hagerup. Field-grown plants were transferred to a cold room kept at 5°C in late autumn and then analysed for the foliage respiration rate in relation to nitrogen and sugar concentration over a period of many weeks. Respiration rate varied 1.6-fold among species at a given time, and decreased with time as long as plants remained dormant. Most of both sources of variation were accounted for by the same linear and positive correlation with total soluble sugar concentration, whereas no relationship with nitrogen concentration was found. The hypothesis presented is that respiration rate correlates with sugar concentration in the dormant season because cellular sugar concentrations are much increased and, thereby, the costs of maintaining concentration gradients. Pinus contorta had a significantly higher respiration rate for a given sugar concentration than any other species, and therefore suffered larger relative losses of sugars when kept at 5°C; possible reasons and consequences of this are discussed in relation to field performance.
Article
Eddy covariance measurements of CO 2 flux, based on four and six week campaigns in Rondôdnia, Brazil, have been used in conjunction with a model to scale up data to a whole year, and thus estimate the carbon balance of the tropical forest ecosystem, and the changes in carbon balance expected from small interannual variations in climatological conditions. One possible source of error in this estimation arises from the difficulty in measuring fluxes under stably stratified meteorological conditions, such as occur frequently at night. Flux may be ‘lost’ because of low velocity advection, caused by nocturnal radiative cooling at sites on raised ground. Such effects may be detected by plotting the net ecosystem flux of CO 2 , F eco is a function of wind speed. If flux is ‘lost’ then F eco is expected to decline with wind speed. In the present data set, this did not occur, and F eco was similar to the nocturnal flux estimated independently from chamber measurements. The model suggests that in 1992/3, the Gross Primary Productivity (GPP) was 203.3 mol C m ⁻² y ⁻¹ and ecosystem respiration was 194.8 mol C m ⁻² y ⁻¹ , giving an ecosystem carbon balance of 8.5 mol C m ⁻² y ⁻¹ , equivalent to a sink of 1.0 ton C ha ⁻¹ y ⁻¹ . However, the sign and magnitude of this figure is very sensitive to temperature, because of the strong influence of temperature on respiration. The model also suggests that the effect of temperature on the net carbon balance is strongly dependent on the partial pressure of CO 2 .
Article
Rates of CO 2 efflux from the floor of a central Siberian Scots pine (Pinus sylvestris) forest were measured using a dynamic closed chamber system and by a eddy covariance system placed 2.5 m above the forest floor. Measurements were undertaken for a full growing season: from early May to early October 1999. Spatial variability as determined by the chamber measurements showed the rate of CO 2 efflux to depend on location, with rates from relatively open areas ("glades") only being about 50% those observed below or around trees. This was despite generally higher temperatures in the glade during the day. A strong relationship between CO 2 efflux rate and root density was observed in early spring, suggesting that lower rates in open areas may have been attributable to fewer roots there. Continuous measurements with the eddy covariance system provided good temporal coverage. This method, however, provided estimates of ground CO 2 efflux rate rates that were about 50% lower than chamber measurements that were undertaken in areas considered to be representative of the forest as a whole. An examination of the seasonal pattern of soil CO 2 efflux rates suggests that much of the variability in CO 2 efflux rate could be accounted for by variations in soil temperature. Nevertheless, there were also some indications that the soil water deficits served to reduce soil CO 2 efflux rates during mid-summer. Overall the sensitivity of CO 2 efflux rate to temperature seems to be greater for this boreal ecosystem than has been the case for most other studies.
Article
The exchange of carbon dioxide (CO 2 ) between the atmosphere and a forest after disturbance by wind throw in the western Russian taiga was investigated between July and October 1998 using the eddy covariance technique. The research area was a regenerating forest (400 m × 1000 m), in which all trees of the preceding generation were uplifted during a storm in 1996. All deadwood had remained on site after the storm and had not been extracted for commercial purposes. Because of the heterogeneity of the terrain, several micrometeorological quality tests were applied. In addition to the eddy covariance measurements, carbon pools of decaying wood in a chronosequence of three different wind throw areas were analysed and the decay rate of coarse woody debris was derived. During daytime, the average CO 2 uptake flux was −3 µmol m ⁻² s ⁻¹ , whereas during night‐time characterised by a well‐mixed atmosphere the rates of release were typically about 6 µmol m ⁻² s ⁻¹ . Suppression of turbulent fluxes was only observed under conditions with very low friction velocity ( u * ≤ 0.08 ms ⁻¹ ). On average, 164 mmol CO 2 m ⁻² d ⁻¹ was released from the wind throw to the atmosphere, giving a total of 14.9 mol CO 2 m ⁻² (180 g CO 2 m ⁻² ) released during the 3‐month study period. The chronosequence of dead woody debris on three different wind throw areas suggested exponential decay with a decay coefficient of −0.04 yr ⁻¹ . From the magnitude of the carbon pools and the decay rate, it is estimated that the decomposition of coarse woody debris accounted for about a third of the total ecosystem respiration at the measurement site. Hence, coarse woody debris had a long‐term influence on the net ecosystem exchange of this wind throw area. From the analysis performed in this work, a conclusion is drawn that it is necessary to include into flux networks the ecosystems that are subject to natural disturbances and that have been widely omitted into considerations of the global carbon budget. The half‐life time of about 17 years for deadwood in the wind throw suggests a fairly long storage of carbon in the ecosystem, and indicates a very different long‐term carbon budget for naturally disturbed vs. commercially managed forests.