Dependence of the aboveground respiration of hinoki cypress (Chamaecyparis obtusa) on tree size

Nighttime respiration was measured at monthly intervals over one year on the aboveground parts of five sample trees in an 8-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) stand, by an enclosed standing-tree method. The respiration rate rose rapidly from early spring to a maximum in June, and decreased abruptly in July and then gradually toward autumn and winter. The seasonal change in the respiration rate was synchronized with stem volume increment rather than with monthly mean air temperature. The respiration rate, r, of individual trees increased with increasing tree dimensions, such as stem volume, v(S), and stem girth at the base of the live crown, G(B). The dependence of respiration rate on tree size was successfully represented by a power function. The r - v(S) dependence was rather stronger than the r - G(B) (2) dependence, especially toward the end of the growing season (from July to September). The observed respiration rate was almost the same as the respiration rate corrected for the monthly mean air temperature. The annual respiration of individual trees was directly proportional to their phytomass or to its increment. Although the annual respiration of individual trees decreased proportionally to the square root of the leaf mass, it decreased abruptly in the range close to the smallest sample tree. Combining the monthly relationship between respiration rate and stem volume with the tree size distribution in the stand, the stand aboveground annual respiration was estimated to be 20.4 Mg CO(2) ha(-1) year(-1) (= 12.5 Mg dry mass ha(-1) year(-1)) for an aboveground biomass of 17.4 Mg ha(-1) with an annual increment of 6.51 Mg ha(-1) year(-1), i.e., the stand aboveground annual respiration amounted to the equivalent of 72% of the biomass or to almost twice the biomass increment.     

Successional changes in live and dead wood carbon stores: implications for net ecosystem productivity

If forests are to be used in CO2 mitigation projects, it is essential to understand and quantify the impacts of disturbance on net ecosystem productivity (NEP; i.e., the change in ecosystem carbon (C) storage with time). We examined the influence of live tree and coarse woody debris (CWD) on NEP during secondary succession based on data collected along a 500-year chronosequence on the Wind River Ranger District, Washington. We developed a simple statistical model of live and dead wood accumulation and decomposition to predict changes in the woody component of NEP, which we call NEP(w). The transition from negative to positive NEP(w), for a series of scenarios in which none to all wood was left after disturbance, occurred between 0 and 57 years after disturbance. The timing of this transition decreased as live-tree growth rates increased, and increased as CWD left after disturbance increased. Maximum and minimum NEP(w) for all scenarios were 3.9 and -14.1 Mg C ha-1 year-1, respectively. Maximum live and total wood C stores of 319 and 393 Mg C ha(-1), respectively, were reached approximately 200 years after disturbance. Decomposition rates (k) of CWD ranged between 0.013 and 0.043 year-1 for individual stands. Regenerating stands took 41 years to attain a mean live wood mass equivalent to the mean mass of CWD left behind after logging, 40 years to equal the mean CWD mass in 500-year-old forest, and more than 150 years to equal the mean total live and dead wood in an old-growth stand. At a rotation age of 80 years, regenerating stands stored approximately half the wood C of the remaining nearby old-growth forests (predominant age 500 years), indicating that conversion of old-growth forests to younger managed forests results in a significant net release of C to the atmosphere.     

Changes in net ecosystem productivity with forest age following clearcutting of a coastal Douglas-fir forest: testing a mathematical model with eddy covariance measurements along a forest chronosequence

We hypothesized that changes in net ecosystem productivity (NEP) during aging of coastal Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands could be explained by (1) changing nutrient uptake caused by different time scales for decomposition of fine, non-woody and coarse woody litter left after harvesting, (2) declines in canopy water status with lengthening of the water uptake pathway during bole and branch growth, and (3) increases in the ratio of autotrophic respiration (R (a)) to gross primary productivity (GPP) with phytomass accumulation. These hypotheses were implemented and tested in the mathematical model ecosys against eddy covariance (EC) measurements of forest CO(2) and energy exchange in a post-clearcut Douglas-fir chronosequence. Hypothesis 1 explained how (a) an initial rise in GPP observed during the first 3 years after clearcutting could be caused by nutrient mineralization from rapid decomposition of fine, non-woody litter with lower C:N ratios (assart effect), (b) a slower rise in GPP during the next 20 years could be caused by immobilization during later decomposition of coarse woody litter, and (c) a rapid rise in GPP between 20 and 40 years after clearcutting could be caused by nutrient mineralization with further decomposition of coarse woody litter and of its decomposition products. During periods (a) and (b), heterotrophic respiration (R (h)) from decomposition of fine and coarse litter greatly exceeded net primary productivity (NPP = GPP - R (a)) so that Douglas-fir stands were large sources of CO(2). During period (c), NPP exceeded R (h) so that these stands became large sinks for CO(2). Hypothesis 2 explained how declines in NPP during later growth in period (c) could be caused by lower hydraulic conductances in taller trees that would force lower canopy water potentials and hence greater sensitivity of stomatal conductances and CO(2) uptake to vapor pressure deficits. Enhanced sensitivity to vapor pressure deficits was also apparent in the EC measurements over the post-clearcut chronosequence. Hypothesis 3 did not contribute to the explanation of forest age effects on NEP.     

Component and whole-system respiration fluxes in northern deciduous forests

We measured component and whole-system respiration fluxes in northern hardwood (Acer saccharum Marsh., Tilia americana L., Fraxinus pennsylvanica Marsh.) and aspen (Populus tremuloides Michx.) forest stands in Price County, northern Wisconsin from 1999 through 2002. Measurements of soil, leaf and stem respiration, stem biomass, leaf area and biomass, and vertical profiles of leaf area were combined with biometric measurements to create site-specific respiration models and to estimate component and whole-system respiration fluxes. Hourly estimates of component respiration were based on site measurements of air, soil and stem temperature, leaf mass, sapwood volume and species composition. We also measured whole-system respiration from an above-canopy eddy flux tower. Measured soil respiration rates varied significantly among sites, but not consistently among dominant species (P < 0.05 and P > 0.1). Annual soil respiration ranged from 8.09 to 11.94 Mg C ha(-1) year(-1). Soil respiration varied linearly with temperature (P < 0.05), but not with soil water content (P > 0.1). Stem respiration rates per unit volume and per unit area differed significantly among species (P < 0.05). Stem respiration per unit volume of sapwood was highest in F. pennsylvanica (up to 300 micro mol m(3) s(-1)) and lowest in T. americana (22 micro mol m(3) s(-1)) when measured at peak summer temperatures (27 to 29 degrees C). In northern hardwood stands, south-side stem temperatures were higher and more variable than north-side temperatures during leaf-off periods, but were not different statistically during leaf-on periods. Cumulative annual stem respiration varied by year and species (P < 0.05) and averaged 1.59 Mg C ha(-1) year(-1). Leaf respiration rates varied significantly among species (P < 0.05). Respiration rates per unit leaf mass measured at 30 degrees C were highest for P. tremuloides (38.8 nmol g(-1) s(-1)), lowest for Ulmus rubra Muhlenb. (13.1 nmol g(-1) s(-1)) and intermediate and similar (30.2 nmol g(-1) s(-1)) for T. americana, F. pennsylvanica and Q. rubra. During the growing season, component respiration estimates were dominated by soil respiration, followed by leaf and then stem respiration. Summed component respiration averaged 11.86 Mg C ha(-1) year(-1). We found strong covariance between whole-ecosystem and summed component respiration measurements, but absolute rates and annual sums differed greatly.     

Carbon sequestration in the trees, products and soils of forest plantations: an analysis using UK examples

A carbon-flow model for managed forest plantations was used to estimate carbon storage in UK plantations differing in Yield Class (growth rate), thinning regime and species characteristics. Time-averaged, total carbon storage (at equilibrium) was generally in the range 40-80 Mg C ha(-1) in trees, 15-25 Mg C ha(-1) in above- and belowground litter, 70-90 Mg C ha(-1) in soil organic matter and 20-40 Mg C ha(-1) in wood products (assuming product lifetime equalled rotation length). The rate of carbon storage during the first rotation in most plantations was in the range 2-5 Mg C ha(-1) year(-1).A sensitivity analysis revealed the following processes to be both uncertain and critical: the fraction of total woody biomass in branches and roots; litter and soil organic matter decomposition rates; and rates of fine root turnover. Other variables, including the time to canopy closure and the possibility of accelerated decomposition after harvest, were less critical. The lifetime of wood products was not critical to total carbon storage because wood products formed only a modest fraction of the total.The average increase in total carbon storage in the tree-soil-product system per unit increase in Yield Class (m(3) ha(-1) year(-1)) for unthinned Picea sitchensis (Bong.) Carr. plantations was 5.6 Mg C ha(-1). Increasing the Yield Class from 6 to 24 m(3) ha(-1) year(-1) increased the rate of carbon storage in the first rotation from 2.5 to 5.6 Mg C ha(-1) year(-1) in unthinned plantations. Thinning reduced total carbon storage in P. sitchensis plantations by about 15%, and is likely to reduce carbon storage in all plantation types.If the objective is to store carbon rapidly in the short term and achieve high carbon storage in the long term, Populus plantations growing on fertile land (2.7 m spacing, 26-year rotations, Yield Class 12) were the best option examined. If the objective is to achieve high carbon storage in the medium term (50 years) without regard to the initial rate of storage, then plantations of conifers of any species with above-average Yield Classes would suffice. In the long term (100 years), broadleaved plantations of oak and beech store as much carbon as conifer plantations. Mini-rotations (10 years) do not achieve a high carbon storage.     

Soil respiration in a mixed temperate forest and its contribution to total ecosystem respiration

Soil respiration (SR) was measured with an infrared gas analyzer in nine plots representative of the heterogeneous vegetation in a mixed coniferous-deciduous forest in the Belgian Campine region. Selected plots included the two most representative overstory species (Pinus sylvestris L. and Quercus robur L.) in combination with the most representative understory species of the forest. A model that includes temperature and water as the main controlling variables was fitted to the data. We found large spatial variability in SR among plots, with typically lower fluxes under the coniferous overstory than under the deciduous overstory (means of 4.8 +/- 0.4 and 8.8 +/- 0.5 Mg C ha(-1) year(-1), respectively). Total annual soil carbon (C) emissions were estimated by weighting fluxes from different types of vegetation according to their relative contribution to the footprint area of the eddy covariance flux measurement. The relative contribution of the two main tree species to the footprint-weighted total SR varied among seasons with the more abundant coniferous overstory contributing the most to total SR during most of the year. Nonetheless, during summer, the contribution of deciduous plots to total SR was disproportionally high because of the more pronounced seasonality of belowground metabolic activity. Net ecosystem carbon dioxide exchange was measured by eddy covariance, and we estimated total ecosystem respiration (TER) with footprint-constrained nighttime fluxes. Mean total annual SR and TER were 6.1 +/- 0.11 and 9.1 +/- 1.15 Mg C ha(-1) year(-1), respectively. The 95% confidence interval of the ratio of annual SR:TER ranged from 0.58 to 0.76, with a mean of 0.67. The contribution of SR to TER tended to vary seasonally, with minimum contributions during summer (less than 50% of TER) and maximum contributions during winter (about 94% of TER).     

Fruit development, not GPP, drives seasonal variation in NPP in a tropical palm plantation

We monitored seasonal variations in net primary production (NPP), estimated by allometric equations from organ dimensions, gross primary production (GPP), estimated by the eddy covariance method, autotrophic respiration (R(a)), estimated by a model, and fruit production in a coconut (Cocos nucifera L.) plantation located in the sub-tropical South Pacific archipelago of Vanuatu. Net primary production of the vegetative compartments of the trees accumulated steadily throughout the year. Fruits accounted for 46% of tree NPP and showed large seasonal variations. On an annual basis, the sum of estimated NPP (16.1 Mg C ha(-1) year(-1)) and R(a) (24.0 Mg C ha(-1) year(-1)) for the ecosystem (coconut trees and herbaceous understory) closely matched GPP (39.0 Mg C ha(-1) year(-1)), suggesting adequate cross-validation of annual C budget methods. However, seasonal variations in NPP + R(a) were smaller than the seasonal variations in GPP, and maximum tree NPP occurred 6 months after the midsummer peak in GPP and solar radiation. We propose that this discrepancy reflects seasonal variation in the allocation of dry mass to carbon reserves and new plant tissue, thus affecting the allometric relationships used for estimating NPP.     

Spatial and Temporal Patterns in Stand Structure,Biomass, Growth,and Mortality in a Monospecific Nothofagus solandrivar. cliffortioides (Hook,f.) Poole Forest in New Zealand

Abstract

In 250 20 m X 20 m permanent plots in the Craigie-burn Range, Canterbury, New Zealand, 1970 stem density was 2,191/ha, basal area was 52.4 m²/ha, and stem biomass was 178.1 Mg/ha. Net production of stemwood (1974-1987) was 2.0 Mg/ha/yr; mortality was 3.5 Mg/ha/yr. By 1987 density had decreased by 30%, basal area by 12%, and stem biomass by 13%.

Stands with many short trees of small mean dbh were common at high elevation, whereas stands with fewer, taller trees with large mean dbh were common at low elevation. Stemwood production and mortality rate were higher in tall stands. Mortality was well distributed among plots, indicating small, frequent canopy openings; stand turnover calculations were 66 year (based on 2.2% annual biomass loss) to 83 year (based on 1.2% annual stemwood production). Larger canopy openings were also evident, but were more infrequent, so stand turnover times due to 'catastrophic' disturbances were in the range of 350-4000 yr. Consequently, the small, high-frequency disturbances blurred effects of larger disturbances on stand structure and also constrained the fluctuation in forest biomass.     

Maintenance and growth respiration of the aboveground parts of young field-grown hinoki cypress (Chamaecyparis obtusa)

Aboveground respiration of five 8-year-old trees of field-grown hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) was nondestructively measured at monthly intervals over 1 year with an enclosed standing tree method. The relationship between monthly specific respiration rate and monthly mean relative growth rate at the individual tree level was described by a linear equation. During the dormant season, respiration was used mainly for maintenance purposes, whereas during the growing season, more than 40% of the respiration was used for growth purposes, i.e., 60 to 70% in May. We conclude that annual maintenance and growth respiration of a tree are directly proportional to the aboveground phytomass and its annual increment, respectively. The maintenance coefficient was estimated to be 0.504 +/- 0.039 (SE) kg kg(-1) year(-1), indicating that the amount respired for maintaining already existing phytomass was equivalent to about half of the existing phytomass. The growth coefficient was estimated to be 0.772 +/- 0.043 (SE) kg kg(-1), indicating that the amount respired for constructing new phytomass was equivalent to about three-fourths of the annual phytomass increment. The annual stand maintenance and growth respiration were, respectively, 8.8 Mg ha(-1) year(-1) for an aboveground biomass of 17.4 Mg ha(-1) and 5.0 Mg ha(-1) year(-1) for an annual stand aboveground biomass increment of 6.5 Mg ha(-1) year(-1). About two-thirds of the total respiration was used to maintain already existing biomass, and about one-third was used to construct new biomass.     

Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.     

Managing forests for wood yield and carbon storage: a theoretical study

Which forest management regimes best achieve the dual objectives of high sustained timber yield and high carbon storage, including the carbon stored in soil and wood products? A mechanistic forest ecosystem simulator, which couples carbon, nitrogen and water (Edinburgh Forest Model), was calibrated to mimic the growth of a pine plantation in a Scottish climate. The model was then run to equilibrium (1) as an undisturbed forest, (2) removing 2.5, 10, 20 or 40% of the woody biomass each year (3) removing 50% of the woody biomass every 20 years, and (4) clear-felling and replanting every 60 years as in conventional plantations in this climate. More carbon was stored in the undisturbed forest (35.2 kg C m(-2)) than in any regime in which wood was harvested. Plantation management gave moderate carbon storage (14.3 kg C m(-2)) and timber yield (15.6 m(3) ha(-1) year(-1)). Notably, annual removal of 10 or 20% of woody biomass per year gave both a high timber yield (25 m(3) ha(-1) year(-1)) and high carbon storage (20 to 24 kg C m(-2)). The efficiency of the latter regimes could be attributed (in the model) to high light interception and net primary productivity, but less evapotranspiration and summer water stress than in the undisturbed forest, high litter input to the soil giving high soil carbon and N(2) fixation, low maintenance respiration and low N leaching owing to soil mineral pool depletion. We conclude that there is no simple inverse relationship between the amount of timber harvested from a forest and the amount of carbon stored. Management regimes that maintain a continuous canopy cover and mimic, to some extent, regular natural forest disturbance are likely to achieve the best combination of high wood yield and carbon storage.     

Net carbon storage in a poplar plantation (POPFACE) after three years of free-air CO2 enrichment

A high-density plantation of three genotypes of Populus was exposed to an elevated concentration of carbon dioxide ([CO(2)]; 550 micromol mol(-1)) from planting through canopy closure using a free-air CO(2) enrichment (FACE) technique. The FACE treatment stimulated gross primary productivity by 22 and 11% in the second and third years, respectively. Partitioning of extra carbon (C) among C pools of different turnover rates is of critical interest; thus, we calculated net ecosystem productivity (NEP) to determine whether elevated atmospheric [CO(2)] will enhance net plantation C storage capacity. Free-air CO(2) enrichment increased net primary productivity (NPP) of all genotypes by 21% in the second year and by 26% in the third year, mainly because of an increase in the size of C pools with relatively slow turnover rates (i.e., wood). In all genotypes in the FACE treatment, more new soil C was added to the total soil C pool compared with the control treatment. However, more old soil C loss was observed in the FACE treatment compared with the control treatment, possibly due to a priming effect from newly incorporated root litter. FACE did not significantly increase NEP, probably as a result of this priming effect.     

Modeling annual production and carbon fluxes of a large managed temperate forest using forest inventories, satellite data and field measurements

We evaluated annual productivity and carbon fluxes over the Fontainebleau forest, a large heterogeneous forest region of 17,000 ha, in terms of species composition, canopy structure, stand age, soil type and water and mineral resources. The model is a physiological process-based forest ecosystem model coupled with an allocation model and a soil model. The simulations were done stand by stand, i.e., 2992 forest management units of simulation. Some input parameters that are spatially variable and to which the model is sensitive were calculated for each stand from forest inventory attributes, a network of 8800 soil pits, satellite data and field measurements. These parameters are: (1) vegetation attributes: species, age, height, maximal leaf area index of the year, aboveground biomass and foliar nitrogen content; and (2) soil attributes: available soil water capacity, soil depth and soil carbon content. Main outputs of the simulations are wood production and carbon fluxes on a daily to yearly basis. Results showed that the forest is a carbon sink, with a net ecosystem exchange of 371 g C m(-2) year(-1). Net primary productivity is estimated at 630 g C m(-2) year(-1) over the entire forest. Reasonably good agreement was found between simulated trunk relative growth rate (2.74%) and regional production estimated from the National Forest Inventory (IFN) (2.52%), as well as between simulated and measured annual wood production at the forest scale (about 71,000 and 68,000 m(3) year(-1), respectively). Results are discussed species by species.     

Interannual variability of net ecosystem production for a broadleaf deciduous forest in Sapporo, northern Japan

To estimate net ecosystem production (NEP), ecosystem respiration (R _E), and gross primary production (GPP), and to elucidate the interannual variability of NEP in a cool temperate broadleaf deciduous forest in Sapporo, northern Japan, we measured net ecosystem exchange (NEE) using an eddy covariance technique with a closed-path infrared gas analyzer from 2000 to 2003. NEP, R _E, and GPP were derived from NEE, and data gaps were filled using empirical regression models with meteorological variables such as photosynthetic active radiation and soil temperature. In general, NEP was positive (CO₂ uptake) from May to September, either positive or negative in October, and negative (CO₂ release) from November to the following April. NEP rapidly increased during leaf expansion in May and reached its maximum in June or July. The four-year averages (±?standard deviation) of annual NEP, GPP, and R _E were 443?±?45, 1,374?±?39, and 931?±?11?g?C?m^?2?year^?1, respectively. The lower annual NEP and GPP in 2000 may have been caused by lower solar radiation in the foliated season. During the foliated season, monthly GPP varied from year to year more than monthly R _E. Variations in the amount of incoming solar radiation may have caused the interannual variations in the monthly GPP. Additionally, in May, the timing of leaf expansion had a large impact on GPP. Variations in GPP affected the interannual variation in NEP at our site. Thus, interannual variation in NEP was affected by the incoming solar radiation and the timing of leaf expansion.     

Modeling topographic effects on net ecosystem productivity of boreal black spruce forests

Current regional estimates of net primary productivity (NPP) of boreal black spruce overlook the large variation in NPP caused by small-scale topographic effects on soil water, temperature and nutrient availability. Topographic effects on black spruce NPP could likely be modeled by simulating the lateral and vertical movement of water, and its effects on soil nutrient transformation and uptake, through three-dimensional watersheds defined by aspects and slopes of their topographic positions. To examine this likelihood, the ecosystem model 'ecosys' was run for 120 years on a transect that included upper- and lower-slope positions and a basin in which a basal water table was set 0.5 m below the soil surface. For the run, we used soil properties and weather conditions recorded at the 115-year-old BOREAS Southern Old Black Spruce site. Short-term model performance was tested by comparing diurnal and annual carbon (C) transfers simulated under 1994 weather conditions during the 115th year of the model run with those measured at this site during 1994 by eddy covariance, surface chambers and allometry. After 115 years, annual spruce NPP simulated at the upper-slope positions was twice that at the basin (350 versus 170 g C m-2), whereas accumulated wood C was almost three times as large (6.8 versus 2.4 kg C m-2). In the model, increases in NPP and wood growth in upper-slope positions were caused by lower soil water contents, higher soil temperatures, and more rapid O2 uptake that accelerated heterotrophic respiration and hence nutrient mineralization and uptake. Modeled differences in wood growth with topographic position were quantitatively consistent with measurements of boreal black spruce at several research sites differing in water table depth. Modeled differences also agreed with differences in wood growth rates derived from allometric measurements at boreal black spruce sites differing in productivity indices as a result of differences in subsurface hydrology. The magnitude of these differences clearly indicates the importance of accounting for subsurface hydrology in regional estimates of boreal forest productivity.     

Inventory and eddy covariance-based estimates of annual carbon sequestration in a Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis> (Bong.) Carr.) forest ecosystem

A comparison was made of annual net ecosystem productivity (NEP) of a closed canopy Sitka spruce forest over 2 years, using either eddy covariance or inventory techniques. Estimates for annual net uptake of carbon (C) by the forest varied between 7.30 and 11.44 t C ha⁻¹ year⁻¹ using ecological inventory (NEP_eco) measures and 7.69–9.44 t C ha⁻¹ year⁻¹ using eddy covariance-based NEP (-NEE) assessments. These differences were not significant due to uncertainties and errors associated with estimates of biomass increment (15–21%) and heterotrophic respiration (12–19%). Carbon-stock change inventory (NEP_ΔC ) values were significantly higher (27–32%), when compared to both NEP_eco- and -NEE-based estimates. Additional analyses of the data obtained from this study, together with published data, suggest that there was a systematic overestimation of NEP_ΔC -based assessments due to unaccounted decomposition processes and uncertainties in the estimation of soil-C stock changes. In contrast, there was no systematic difference between NEP_eco and eddy covariance assessments across a wide range of forest types and geographical locations.     

Changes in the relationship between tree size and aboveground respiration in field-grown hinoki cypress (Chamaecyparis obtusa) trees over three years

Respiration measurements of aerial parts of 18-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) trees were made under field conditions over three years to study changing relationships with tree age between respiration and phytomass, phytomass increment, and leaf mass. The relationship between annual respiration (r(a)) and phytomass (w(T)) was approximated by a proportional function (r(a) = aw(T)), where the proportional constant (a) decreased year by year. The effect of time on the relationship between annual respiration and phytomass of each sample tree was fitted by a power function. Respiration of the tree suppressed by the canopy decreased year by year, but respiration of the other trees increased slightly with age. The relationship between annual respiration and leaf mass was also approximated by a generalized power function. Excluding the suppressed tree, the relationship between annual respiration (r(a)) and the annual increment of aboveground phytomass (Deltaw(T)) was described by a proportional function (r(a) = 2.27Deltaw(T)), where the proportional constant, 2.27, was independent of sample tree and year, indicating that about 2.3 times of the annual aboveground phytomass increment equivalent was respired annually. For any tree, the time constant relationships between annual respiration and leaf mass and phytomass increment for different-sized trees were similar to the corresponding time continuum relationships. In contrast, the time continuum relationship between annual respiration and phytomass differed from the time constant relationship, indicating that respiration of less active woody tissue contributed significantly to aboveground respiration. Based on the relationship between tree size and annual respiration, annual aboveground stand respiration was estimated to be 25.0, 26.9, and 25.8 Mg(dm) ha(-1) year(-1) for the three consecutive years, respectively, and the corresponding aboveground stand biomass was 60.0, 69.0, and 76.8 Mg(dm) ha(-1).     

Seasonal and interannual variation in net ecosystem production of an evergreen needleleaf forest in Japan

Carbon dioxide (CO₂) flux was measured above the forest at the Fujiyoshida site on the northern slope of Mount Fuji in Japan in 2000?C2008 using an eddy covariance technique. The forest mainly consists of Japanese red pine (Pinus densiflora) and Japanese holly (Ilex pedunculosa). The 9-year average of monthly mean net ecosystem production (NEP) ranged from ?0.1?g?C?m^?2?day^?1 in January to 2.5?g?C?m^?2?day^?1 in May. The maximum net uptake was observed in May, although gross primary production (GPP) was highest in July. Variation in the leaf amount did not notably affect seasonal variation in GPP. This site was characterized by carbon uptake even in winter, if the meteorological conditions were conducive for photosynthesis and a resulting long period of carbon uptake. The 9-year averages of annual NEP, GPP, and ecosystem respiration (RE) were 388, 1,802, and 1,413?g?C?m^?2?year^?1, respectively. The annual NEP was lowest in 2003 and highest in 2004 over the 9?years. Year-to-year variability of NEP mainly depended on air temperature and photosynthetically active radiation in summer, and the dependence of the deviation of annual NEP on that of GPP was greater than that of RE. Long-term observational data indicated that the carbon uptake ability at the study site was at a moderate level in comparison with other temperate humid evergreen forests around the world. These data also indicated that the site had a high carbon uptake ability compared with other deciduous forests in Japan because of the duration of carbon uptake.     

Carbon stocks and soil respiration rates during deforestation, grassland use and subsequent Norway spruce afforestation in the Southern Alps, Italy

Changes in carbon stocks during deforestation, reforestation and afforestation play an important role in the global carbon cycle. Cultivation of forest lands leads to substantial losses in both biomass and soil carbon, whereas forest regrowth is considered to be a significant carbon sink. We examined below- and aboveground carbon stocks along a chronosequence of Norway spruce (Picea abies (L.) Karst.) stands (0-62 years old) regenerating on abandoned meadows in the Southern Alps. A 130-year-old mixed coniferous Norway spruce-white fir (Abies alba Mill.) forest, managed by selection cutting, was used as an undisturbed control. Deforestation about 260 years ago led to carbon losses of 53 Mg C ha(-1) from the organic layer and 12 Mg C ha(-1) from the upper mineral horizons (Ah, E). During the next 200 years of grassland use, the new Ah horizon sequestered 29 Mg C ha(-1). After the abandonment of these meadows, carbon stocks in tree stems increased exponentially during natural forest succession, levelling off at about 190 Mg C ha(-1) in the 62-year-old Norway spruce and the 130-year-old Norway spruce-white fir stands. In contrast, carbon stocks in the organic soil layer increased linearly with stand age. During the first 62 years, carbon accumulated at a rate of 0.36 Mg C ha(-1) year(-1) in the organic soil layer. No clear trend with stand age was observed for the carbon stocks in the Ah horizon. Soil respiration rates were similar for all forest stands independently of organic layer thickness or carbon stocks, but the highest rates were observed in the cultivated meadow. Thus, increasing litter inputs by forest vegetation compared with the meadow, and constantly low decomposition rates of coniferous litter were probably responsible for continuous soil carbon sequestration during forest succession. Carbon accumulation in woody biomass seemed to slow down after 60 to 80 years, but continued in the organic soil layer. We conclude that, under present climatic conditions, forest soils act as more persistent carbon sinks than vegetation that will be harvested, releasing the carbon sequestered during tree growth.     

Land-use changes alter CO2 flux patterns of a tall-grass Andropogon field and a savanna-woodland continuum in the Orinoco lowlands

Land use changes in the savannas of the Orinoco lowlands have resulted in a mosaic of vegetation. To elucidate how these changes have affected carbon exchanges with the atmosphere, we measured CO2 fluxes by eddy covariance and soil CO2 efflux systems along a disturbance gradient beginning with a cultivated tall-grass Andropogon field (S1) and extending over three savanna sites with increasing woody cover growing above native herbaceous vegetation. The savanna sites included a herbaceous savanna (S2), a tree savanna (S3) and a woodland savanna (S4). During the wet season, maximum diurnal net ecosystem exchange (NEE) over the S1-S4 sites was 6.6-9.3, 6.6-7.9, 10.6-11.3 and 9.3-10.6 micromol m(-2) s(-1), respectively. The rate of CO2 uptake over S1 was lower than that for C4 grasses elsewhere because of pasture degradation. Soil respiration and temperature were exponentially related when soil water content (theta) was above 0.083 m(3) m(-3); however, soil respiration declined markedly as theta decreased from 0.083-0.090 to 0.033-0.056 m(3) m(-3). There were bursts of CO2 emission when dry soils were rewetted by rainfall. During the wet season, all sites constituted carbon sinks with maximum net daily ecosystem production (NEP) of 2.1, 1.7, 2.1 and 2.1 g C m(-2) day(-1), respectively. During the dry season, the savanna sites (S2-S4) became carbon sources with maximum emission fluxes of -0.5, -1.4 and -1.6 g C m(-2) day(-1), respectively, whereas the tall-grass field (S1) remained a carbon sink with a maximum NEP of 0.3 g C m(-2) day(-1) at the end of the season. For all measurement periods, annual NEP of sites S1-S4 was 366, 6, 116 and 139 g C m(-2), respectively. Comparisons of carbon source/sink dynamics across a wide range of savannas indicate that savanna carbon budgets can change in sign and magnitude. On an annual basis, gross primary production over the S1-S4 stands was 797, 803, 136 and 1230 g C m(-2), respectively. Net primary productivity (NPP) of the S1-S4 stands, calculated from eddy covariance measurements as the daily sum of NEE and day and night heterotrophic respiration was 498, 169, 181 and 402 g C m-2 year-1, respectively. These values were slightly higher than NPP based on harvest measurements (432, 162, 176 and 386 g C m(-2) year(-1), respectively), presumably because fine roots were incompletely harvested. Soil water content limited carbon uptake at all sites, and water-use efficiency (WUE) was related to rainfall dynamics. During the dry season, all sites except the cultivated tall-grass Andropogon field (S1) had a negative WUE. Although our results are specific to the Orinoco vegetational mosaic, the effects of land-use practices on the controls and physiological functions of the studied ecosystems may be generalized to other savannas.