Elevated CO2 affects photosynthetic responses in canopy pine and subcanopy deciduous trees over 10 years: a synthesis from Duke FACE

Leaf responses to elevated atmospheric CO₂ concentration (C_a) are central to models of forest CO₂ exchange with the atmosphere and constrain the magnitude of the future carbon sink. Estimating the magnitude of primary productivity enhancement of forests in elevated C_a requires an understanding of how photosynthesis is regulated by diffusional and biochemical components and up‐scaled to entire canopies. To test the sensitivity of leaf photosynthesis and stomatal conductance to elevated C_a in time and space, we compiled a comprehensive dataset measured over 10 years for a temperate pine forest of Pinus taeda, but also including deciduous species, primarily Liquidambar styraciflua. We combined over one thousand controlled‐response curves of photosynthesis as a function of environmental drivers (light, air C_a and temperature) measured at canopy heights up to 20 m over 11 years (1996–2006) to generate parameterizations for leaf‐scale models for the Duke free‐air CO₂ enrichment (FACE) experiment. The enhancement of leaf net photosynthesis (A_net) in P. taeda by elevated C_a of +200 μmol mol^?1 was 67% for current‐year needles in the upper crown in summer conditions over 10 years. Photosynthetic enhancement of P. taeda at the leaf‐scale increased by two‐fold from the driest to wettest growing seasons. Current‐year pine foliage A_net was sensitive to temporal variation, whereas previous‐year foliage A_net was less responsive and overall showed less enhancement (+30%). Photosynthetic downregulation in overwintering upper canopy pine needles was small at average leaf N (N_area), but statistically significant. In contrast, co‐dominant and subcanopy L. styraciflua trees showed A_net enhancement of 62% and no A_net–N_area adjustments. Various understory deciduous tree species showed an average A_net enhancement of 42%. Differences in photosynthetic responses between overwintering pine needles and subcanopy deciduous leaves suggest that increased C_a has the potential to enhance the mixed‐species composition of planted pine stands and, by extension, naturally regenerating pine‐dominated stands.     

Global patterns and predictors of stem CO2 efflux in forest ecosystems

Stem CO₂ efflux (E_S) plays an important role in the carbon balance of forest ecosystems. However, its primary controls at the global scale are poorly understood and observation‐based global estimates are lacking. We synthesized data from 121 published studies across global forest ecosystems and examined the relationships between annual E_S and biotic and abiotic factors at individual, biome, and global scales, and developed a global gridded estimate of annual E_S. We tested the following hypotheses: (1) Leaf area index (LAI) will be highly correlated with annual E_S at biome and global scales; (2) there will be parallel patterns in stem and root CO₂ effluxes (R_A) in all forests; (3) annual E_S will decline with forest age; and (4) LAI coupled with mean annual temperature (MAT) and mean annual precipitation (MAP) will be sufficient to predict annual E_S across forests in different regions. Positive linear relationships were found between E_S and LAI, as well as gross primary production (GPP), net primary production (NPP), wood NPP, soil CO₂ efflux (R_S), and R_A. Annual E_S was correlated with R_A in temperate forests after controlling for GPP and MAT, suggesting other additional factors contributed to the relationship. Annual E_S tended to decrease with stand age. Leaf area index, MAT and MAP, predicted 74% of variation in E_S at global scales. Our statistical model estimated a global annual E_S of 6.7 ± 1.1 Pg C yr⁻¹ over the period of 2000–2012 with little interannual variability. Modeled mean annual E_S was 71 ± 43, 270 ± 103, and 420 ± 134 g C m² yr⁻¹ for boreal, temperate, and tropical forests, respectively. We recommend that future studies report E_S at a standardized constant temperature, incorporate more manipulative treatments, such as fertilization and drought, and whenever possible, simultaneously measure both aboveground and belowground CO₂ fluxes.     

Chlorophyll fluorescence tracks seasonal variations of photosynthesis from leaf to canopy in a temperate forest

Accurate estimation of terrestrial gross primary productivity (GPP) remains a challenge despite its importance in the global carbon cycle. Chlorophyll fluorescence (ChlF) has been recently adopted to understand photosynthesis and its response to the environment, particularly with remote sensing data. However, it remains unclear how ChlF and photosynthesis are linked at different spatial scales across the growing season. We examined seasonal relationships between ChlF and photosynthesis at the leaf, canopy, and ecosystem scales and explored how leaf‐level ChlF was linked with canopy‐scale solar‐induced chlorophyll fluorescence (SIF) in a temperate deciduous forest at Harvard Forest, Massachusetts, USA. Our results show that ChlF captured the seasonal variations of photosynthesis with significant linear relationships between ChlF and photosynthesis across the growing season over different spatial scales (R² = 0.73, 0.77, and 0.86 at leaf, canopy, and satellite scales, respectively; P < 0.0001). We developed a model to estimate GPP from the tower‐based measurement of SIF and leaf‐level ChlF parameters. The estimation of GPP from this model agreed well with flux tower observations of GPP (R² = 0.68; P < 0.0001), demonstrating the potential of SIF for modeling GPP. At the leaf scale, we found that leaf F_q’/F_m’, the fraction of absorbed photons that are used for photochemistry for a light‐adapted measurement from a pulse amplitude modulation fluorometer, was the best leaf fluorescence parameter to correlate with canopy SIF yield (SIF/APAR, R² = 0.79; P < 0.0001). We also found that canopy SIF and SIF‐derived GPP (GPP_SIF) were strongly correlated to leaf‐level biochemistry and canopy structure, including chlorophyll content (R² = 0.65 for canopy GPP_SIF and chlorophyll content; P < 0.0001), leaf area index (LAI) (R² = 0.35 for canopy GPP_SIF and LAI; P < 0.0001), and normalized difference vegetation index (NDVI) (R² = 0.36 for canopy GPP_SIF and NDVI; P < 0.0001). Our results suggest that ChlF can be a powerful tool to track photosynthetic rates at leaf, canopy, and ecosystem scales.     

Determinants of soil organic carbon sequestration and its contribution to ecosystem carbon sinks of planted forests

The area of forest established through afforestation/reforestation has been increasing on a global scale, which is particularly important as these planted forests attenuate climate change by sequestering carbon. However, the determinants of soil organic carbon (SOC) sequestration and their contribution to the ecosystem carbon sink of planted forests remain uncertain. By using globally distributed data extracted from 154 peer‐reviewed publications and a total of 355 sampling points, we investigated above‐ground biomass carbon (ABC) sequestration and SOC sequestration across three different climatic zones (tropical, warm temperate, and cold temperate) through correlation analysis, regression models, and structural equation modeling (SEM). We found that the proportion of SOC sequestration in the ecosystem C sequestration averaged 14.1% globally, being the highest (27.0%) in the warm temperate and the lowest (10.7%) in the tropical climatic zones. The proportion was mainly affected by latitude. The sink rate of ABC (R_ABC) in tropical climates (2.48 Mg C ha^?1 year^?1) and the sink rate of SOC (R_SOC) in warm temperate climates (0.96 Mg C ha^?1 year^?1) were higher than other climatic zones. The main determinants of R_SOC were the number of frost‐free days, latitude, mean annual precipitation (MAP), and SOC density (SOCD) at the initial observation; however, these variables depended on the climatic zone. According to the SEM, frost‐free period, mean annual temperature (MAT) and MAP are the dominant driving factors affecting R_SOC in Chinese plantations. MAT has a positive effect on R_SOC, and global warming may increase R_SOC of temperate plantations in China. Our findings highlight the determinants of SOC sequestration and quantitatively reveal the substantial global contribution of SOC sequestration to ecosystem carbon sink provided by planted forests. Our results help managers identify and control key factors to increase carbon sequestration in forest ecosystems.     

The carbon balance in natural and disturbed forests of the southern taiga in central Siberia

Abstract. We evaluated the balance of production and decomposition in natural ecosystems of Pinus sylvestris, Larix sibirica and Betula pendula in the southern boreal forests of central Siberia, using the Yenisei transect. We also investigated whether anthropogenic disturbances (logging, fire and recreation pressure) influence the carbon budget. Pinus and Larix stands up to age class VI act as a net sink for atmospheric carbon. Mineralization rates in young Betula forests exceed rates of uptake via photosynthesis assimilation. Old‐growth stands of all three forest types are CO₂ sources to the atmosphere. The prevalence of old‐growth Larix in the southern taiga suggests that Larix stands are a net source of CO₂. The CO₂ flux to the atmosphere exceeds the uptake of atmospheric carbon via photosynthesis by 0.23 t C.ha^‐1.yr^‐1 (47%). Betula and Pinus forests are net sinks, as photosynthesis exceeds respiration by 13% and 16% respectively. The total carbon flux from Pinus, Larix and Betula ecosystems to the atmosphere is 10 387 thousand tons C.yr^‐1. Net Primary Production (0.935 t‐C.ha^‐1) exceeds carbon release from decomposition of labile and mobile soil organic matter (Rh) by 767 thousand tons C (0.064 t‐C.ha^‐1), so that these forests are net C‐sinks. The emissions due to decomposition of slash (101 thousand tons C; 1.0%) and from fires (0.21%) are very small. The carbon balance of human‐disturbed forests is significantly different. A sharp decrease in biomass stored in Pinus and Betula ecosystems leads to decreased production. As a result, the labile organic matter pool decreased by 6–8 times; course plant residues with a low decomposition rate thus dominate this pool. Annual carbon emissions to the atmosphere from these ecosystems are determined primarily by decomposing fresh litterfall. This source comprises 40–79% of the emissions from disturbed forests compared to only 13–28% in undisturbed forests. The ratio of emissions to production (NPP) is 20–30% in disturbed and 52–76% in undisturbed forests.     

Trends and methodological impacts in soil CO2 efflux partitioning: A metaanalytical review

Partitioning soil carbon dioxide (CO₂) efflux (R_S) into autotrophic (R_A; including plant roots and closely associated organisms) and heterotrophic (R_H) components has received considerable attention, as differential responses of these components to environmental change have profound implications for the soil and ecosystem C balance. The increasing number of partitioning studies allows a more detailed analysis of experimental constraints than was previously possible. We present results of an exhaustive literature search of partitioning studies and analyse global trends in flux partitioning between biomes and ecosystem types by means of a metaanalysis. Across all data, an overall decline in the R_H/R_S ratio for increasing annual R_S fluxes emerged. For forest ecosystems, boreal coniferous sites showed significantly higher (P<0.05) R_H/R_S ratios than temperate sites, while both temperate or tropical deciduous forests did not differ in ratios from any of the other forest types. While chronosequence studies report consistent declines in the R_H/R_S ratio with age, no difference could be detected for different age groups in the global data set. Different methodologies showed generally good agreement if the range of R_S under which they had been measured was considered, with the exception of studies estimating R_H by means of root mass regressions against R_S, which resulted in consistently lower R_H/R_S estimates out of all methods included. Additionally, the time step over which fluxes were partitioned did not affect R_H/R_S ratios consistently. To put results into context, we review the most common techniques and point out the likely sources of errors associated with them. In order to improve soil CO₂ efflux partitioning in future experiments, we include methodological recommendations, and also highlight the potential interactions between soil components that may be overlooked as a consequence of the partitioning process itself.     

Estimates of soil respiration and net primary production of three forests at different succession stages in South China

Soil respiration (heterotropic and autotropic respiration, R_g) and aboveground litter fall carbon were measured at three forests at different succession (early, middle and advanced) stages in Dinghushan Biosphere Reserve, Southern China. It was found that the soil respiration increases exponentially with soil temperature at 5 cm depth (T_s) according to the relation R_g=a exp(bT_s), and the more advanced forest community during succession has a higher value of a because of higher litter carbon input than the forests at early or middle succession stages. It was also found that the monthly soil respiration is linearly correlated with the aboveground litter carbon input of the previous month. Using measurements of aboveground litter and soil respiration, the net primary productions (NPPs) of three forests were estimated using nonlinear inversion. They are 475, 678 and 1148 g C m^?2 yr^?1 for the Masson pine forest (MPF), coniferous and broad‐leaf mixed forest (MF) and subtropical monsoon evergreen broad‐leaf forest (MEBF), respectively, in year 2003/2004, of which 54%, 37% and 62% are belowground NPP for those three respective forests if no change in live plant biomass is assumed. After taking account of the decrease in live plant biomass, we estimated the NPP of the subtropical MEBF is 970 g C m^?2 yr^?1 in year 2003/2004. Total amount of carbon allocated below ground for plant roots is 388 g C m^?2 yr^?1 for the MPF, 504 g C m^?2 yr^?1 for the coniferous and broad‐leaf MF and 1254 g C m^?2 yr^?1 for the subtropical MEBF in 2003/2004. Our results support the hypothesis that the amount of carbon allocation belowground increases during forest succession.     

Higher than expected CO2 fertilization inferred from leaf to global observations

Several lines of evidence point to an increase in the activity of the terrestrial biosphere over recent decades, impacting the global net land carbon sink (NLS) and its control on the growth of atmospheric carbon dioxide (c_a). Global terrestrial gross primary production (GPP)—the rate of carbon fixation by photosynthesis—is estimated to have risen by (31 ± 5)% since 1900, but the relative contributions of different putative drivers to this increase are not well known. Here we identify the rising atmospheric CO₂ concentration as the dominant driver. We reconcile leaf‐level and global atmospheric constraints on trends in modeled biospheric activity to reveal a global CO₂ fertilization effect on photosynthesis of 30% since 1900, or 47% for a doubling of c_a above the pre‐industrial level. Our historic value is nearly twice as high as current estimates (17 ± 4)% that do not use the full range of available constraints. Consequently, under a future low‐emission scenario, we project a land carbon sink (174 PgC, 2006–2099) that is 57 PgC larger than if a lower CO₂ fertilization effect comparable with current estimates is assumed. These findings suggest a larger beneficial role of the land carbon sink in modulating future excess anthropogenic CO₂ consistent with the target of the Paris Agreement to stay below 2°C warming, and underscore the importance of preserving terrestrial carbon sinks.     

Carbon storage and fluxes in ponderosa pine forests at different developmental stages

We compared carbon storage and fluxes in young and old ponderosa pine stands in Oregon, including plant and soil storage, net primary productivity, respiration fluxes, eddy flux estimates of net ecosystem exchange (NEE), and Biome‐BGC simulations of fluxes. The young forest (Y site) was previously an old‐growth ponderosa pine forest that had been clearcut in 1978, and the old forest (O site), which has never been logged, consists of two primary age classes (50 and 250 years old). Total ecosystem carbon content (vegetation, detritus and soil) of the O forest was about twice that of the Y site (21 vs. 10 kg C m^?2 ground), and significantly more of the total is stored in living vegetation at the O site (61% vs. 15%). Ecosystem respiration (R_e) was higher at the O site (1014 vs. 835 g C m^?2 year^?1), and it was largely from soils at both sites (77% of R_e). The biological data show that above‐ground net primary productivity (ANPP), NPP and net ecosystem production (NEP) were greater at the O site than the Y site. Monte Carlo estimates of NEP show that the young site is a source of CO₂ to the atmosphere, and is significantly lower than NEP(O) by c. 100 g C m^?2 year^?1. Eddy covariance measurements also show that the O site was a stronger sink for CO₂ than the Y site. Across a 15‐km swath in the region, ANPP ranged from 76 g C m^?2 year^?1 at the Y site to 236 g C m^?2 year^?1 (overall mean 158 ± 14 g C m^?2 year^?1). The lowest ANPP values were for the youngest and oldest stands, but there was a large range of ANPP for mature stands. Carbon, water and nitrogen cycle simulations with the Biome‐BGC model suggest that disturbance type and frequency, time since disturbance, age‐dependent changes in below‐ground allocation, and increasing atmospheric concentration of CO₂ all exert significant control on the net ecosystem exchange of carbon at the two sites. Model estimates of major carbon flux components agree with budget‐based observations to within ± 20%, with larger differences for NEP and for several storage terms. Simulations showed the period of regrowth required to replace carbon lost during and after a stand‐replacing fire (O) or a clearcut (Y) to be between 50 and 100 years. In both cases, simulations showed a shift from net carbon source to net sink (on an annual basis) 10–20 years after disturbance. These results suggest that the net ecosystem production of young stands may be low because heterotrophic respiration, particularly from soils, is higher than the NPP of the regrowth. The amount of carbon stored in long‐term pools (biomass and soils) in addition to short‐term fluxes has important implications for management of forests in the Pacific North‐west for carbon sequestration.     

Recovery of ponderosa pine ecosystem carbon and water fluxes from thinning and stand‐replacing fire

Carbon uptake by forests is a major sink in the global carbon cycle, helping buffer the rising concentration of CO₂ in the atmosphere, yet the potential for future carbon uptake by forests is uncertain. Climate warming and drought can reduce forest carbon uptake by reducing photosynthesis, increasing respiration, and by increasing the frequency and intensity of wildfires, leading to large releases of stored carbon. Five years of eddy covariance measurements in a ponderosa pine (Pinus ponderosa)‐dominated ecosystem in northern Arizona showed that an intense wildfire that converted forest into sparse grassland shifted site carbon balance from sink to source for at least 15 years after burning. In contrast, recovery of carbon sink strength after thinning, a management practice used to reduce the likelihood of intense wildfires, was rapid. Comparisons between an undisturbed‐control site and an experimentally thinned site showed that thinning reduced carbon sink strength only for the first two posttreatment years. In the third and fourth posttreatment years, annual carbon sink strength of the thinned site was higher than the undisturbed site because thinning reduced aridity and drought limitation to carbon uptake. As a result, annual maximum gross primary production occurred when temperature was 3 °C higher at the thinned site compared with the undisturbed site. The severe fire consistently reduced annual evapotranspiration (range of 12–30%), whereas effects of thinning were smaller and transient, and could not be detected in the fourth year after thinning. Our results show large and persistent effects of intense fire and minor and short‐lived effects of thinning on southwestern ponderosa pine ecosystem carbon and water exchanges.     

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

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

Seasonal changes in canopy photosynthesis and foliage respiration in a Rhizophora stylosa stand at the northern limit of its natural distribution

Gross photosynthesis and respiration rates of leaves at different canopy heights in a Rhizophora stylosa Griff. stand were measured monthly over 1 year at Manko Wetland, Okinawa Island, Japan, which is the northern limit of its distribution. The light-saturated net photosynthesis rate for the leaves at the top of the canopy showed a maximum value of 17 μmol CO₂ m⁻² s⁻¹ in warm season and a minimum value of 6 μmol CO₂ m⁻² s⁻¹ in cold season. The light-saturated gross photosynthesis and dark respiration rates of the leaves existing at the top of the canopy were 2−7 times and 3–16 times, respectively, those of leaves at the bottom of the canopy throughout the year. The light compensation point of leaves showed maximum and minimum peaks in warm season and cold season, respectively. The annual canopy gross photosynthesis, foliage respiration, and surplus production were estimated as 117, 49, and 68 t CO₂ ha⁻¹ year⁻¹, respectively. The energy efficiency of the annual canopy gross photosynthesis was 2.5%. The gross primary production GPP fell near the regression curve of GPP on the product of leaf area index and warmth index, the regression curve which was established for forests in the Western Pacific with humid climates.     

Interactions between CO2 enhancement and N addition on net primary productivity and water‐use efficiency in a mesocosm with multiple subtropical tree species

Carbon dioxide (CO₂) enhancement (eCO₂) and N addition (aN) have been shown to increase net primary production (NPP) and to affect water‐use efficiency (WUE) for many temperate ecosystems, but few studies have been made on subtropical tree species. This study compared the responses of NPP and WUE from a mesocosm composing five subtropical tree species to eCO₂ (700 ppm), aN (10 g N m^?2 yr^?1) and eCO₂ × aN using open‐top chambers. Our results showed that mean annual ecosystem NPP did not changed significantly under eCO₂, increased by 56% under aN and 64% under eCO₂ × aN. Ecosystem WUE increased by 14%, 55%, and 61% under eCO₂, aN and eCO₂ × aN, respectively. We found that the observed responses of ecosystem WUE were largely driven by the responses of ecosystem NPP. Statistical analysis showed that there was no significant interactions between eCO₂ and aN on ecosystem NPP (P = 0.731) or WUE (P = 0.442). Our results showed that increasing N deposition was likely to have much stronger effects on ecosystem NPP and WUE than increasing CO₂ concentration for the subtropical forests. However, different tree species responded quite differently. aN significantly increased annual NPP of the fast‐growing species (Schima superba). Nitrogen‐fixing species (Ormosia pinnata) grew significantly faster only under eCO₂ × aN. eCO₂ had no effects on annual NPP of those two species but significantly increased annual NPP of other two species (Castanopsis hystrix and Acmena acuminatissima). Differential responses of the NPP among different tree species to eCO₂ and aN will likely have significant implications on the species composition of subtropical forests under future global change.     

Carbon budgeting in plant–soil mesocosms under elevated CO2: locally missing carbon?

Studies have suggested that more carbon is fixed due to a large increase in photosynthesis in plant–soil systems exposed to elevated CO₂ than could subsequently be found in plant biomass and soils –‐ the locally missing carbon phenomenon. To further understand this phenomenon, an experiment was carried out using EcoCELLs which are open‐flow, mass‐balance systems at the mesocosm scale. Naturally occurring ¹³C tracers were also used to separately measure plant‐derived carbon and soil‐derived carbon. The experiment included two EcoCELLs, one under ambient atmospheric CO₂ and the other under elevated CO₂ (ambient plus 350 μL L^{? 1}). By matching carbon fluxes with carbon pools, the issue of locally missing carbon was investigated. Flux‐based net primary production (NPP_f) was similar to pool‐based primary production (NPP_p) under ambient CO₂, and the discrepancy between the two carbon budgets (12 g C m^{? 2}, or 4% of NPP_f) was less than measurement errors. Therefore, virtually all carbon entering the system under ambient CO₂ was accounted for at the end of the experiment. Under elevated CO₂, however, the amount of NPP_f was much higher than NPP_p, resulting in missing carbon of approximately 80 g C m^{? 2} or 19% of NPP_f which was much higher than measurement errors. This was additional to the 96% increase in rhizosphere respiration and the 50% increase in root growth, two important components of locally missing carbon. The mystery of locally missing carbon under elevated CO₂ remains to be further investigated. Volatile organic carbon, carbon loss due to root washing, and measurement errors are discussed as some of the potential contributing factors.     

Soil respiration in six temperate forests in China

Scaling soil respiration (R_S), the major CO₂ source to the atmosphere from terrestrial ecosystems, from chamber‐based measurements to ecosystems requires studies on variations and correlations of R_S from various biomes and across geographic regions. However, few studies on R_S are available for Chinese temperate forest despite the importance of this forest in the national and global carbon budgets. In this study, we conducted 18‐month R_S measurements during 2004–2005 in six temperate forest types, representing the typical secondary forest ecosystems across various site conditions in northeastern China: Mongolian oak (Quercus mongolica Fisch.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), mixed deciduous (no dominant tree species), hardwood (dominated by Fraxinus mandshurica Rupr., Juglans mandshurica Maxim., and Phellodendron amurense Rupr.) forests, Korean pine (Pinus koraiensis Sieb. et Zucc.) and Dahurian larch (Larix gmelinii Rupr.) plantations. Our specific objectives were to: (1) explore relationships of R_S against soil temperature and water content for the six forest ecosystems, (2) quantify annual soil surface CO₂ flux and its relations to belowground carbon storage, (3) examine seasonal variations in R_S and related environmental factors, and (4) quantify among‐ and within‐ecosystem variations in R_S. The R_S was positively correlated to soil temperature in all forest types, and was significantly influenced by the interactions of soil temperature and water content in the pine, larch, and mixed deciduous forests. The sensitivity of R_S to soil temperature at 10 cm depth (Q₁₀) ranged from 2.61 in the oak forest to 3.75 in the aspen‐birch forests. The Q₁₀ tended to increase with soil water content until reaching a threshold, and then decline. The annual R_S for the larch, pine, hardwood, oak, mixed deciduous, and aspen‐birch forests averaged 403, 514, 781, 785, 786, and 813 g C m^?2 yr^?1, respectively. The annual R_S of the broadleaved forests was 72% greater than that of the coniferous forests. The annual R_S was positively correlated to soil organic carbon (SOC) concentration at O horizon (R²=0.868) and total biomass of roots <0.5 cm in diameter (R²=0.748). The coefficient of variation (CV) of R_S among forest types averaged 25% across the 18‐month measurements. The CV of R_S within plots varied from 20% to 27%, significantly (P<0.001) greater than those among plots (9–15%), indicating the importance of the fine‐scaled heterogeneity in R_S. This study emphasized that variations in soil respiration and potential sampling bias should be appropriately tackled for accurate soil CO₂ flux estimates.     

Successional changes in soil nitrogen availability,non-symbiotic nitrogen fixation and carbon/nitrogen ratios in southern Chilean forest ecosystems

Vast areas of southern Chile are now covered by second-growth forests because of fire and logging. To study successional patterns after moderate-intensity, anthropogenic fire disturbance, we assessed differences in soil properties and N fluxes across a chronosequence of seven successional stands (2–130 years old). We examined current predictions of successional theory concerning changes in the N cycle in forest ecosystems. Seasonal fluctuations of net N mineralization (N_min) in surface soil and N availability (N_a; N_a=NH ₄ ⁺ –N+NO ₃ ^– –N) in upper and deep soil horizons were positively correlated with monthly precipitation. In accordance with theoretical predictions, stand age was positively, but weakly related to both N_a (r ²=0.282, P<0.001) and total N (N_tot; r ²=0.192, P<0.01), and negatively related to soil C/N ratios (r ²=0.187, P<0.01) in surface soils. A weak linear increase in soil N_min (upper plus deep soil horizons) was found across the chronosequence (r ²=0.124, P<0.022). N_min occurred at modest rates in early successional stands, suggesting that soil disturbance did not impair microbial processes. The relationship between N fixation (N_fix) in the litter layer and stand age best fitted a quadratic model (r ²=0.228, P<0.01). In contrast to documented successional trends for most temperate, tropical and Mediterranean forests, non-symbiotic N_fix in the litter layer is a steady N input to unpolluted southern temperate forests during mid and late succession, which may compensate for hydrological losses of organic N from old-growth ecosystems.     

Strong thermal acclimation of photosynthesis in tropical and temperate wet‐forest tree species: the importance of altered Rubisco content

Understanding of the extent of acclimation of light‐saturated net photosynthesis (A_n) to temperature (T), and associated underlying mechanisms, remains limited. This is a key knowledge gap given the importance of thermal acclimation for plant functioning, both under current and future higher temperatures, limiting the accuracy and realism of Earth system model (ESM) predictions. Given this, we analysed and modelled T‐dependent changes in photosynthetic capacity in 10 wet‐forest tree species: six from temperate forests and four from tropical forests. Temperate and tropical species were each acclimated to three daytime growth temperatures (T_growth): temperate – 15, 20 and 25 °C; tropical – 25, 30 and 35 °C. CO₂ response curves of A_n were used to model maximal rates of RuBP (ribulose‐1,5‐bisphosphate) carboxylation (V_cmax) and electron transport (J_max) at each treatment's respective T_growth and at a common measurement T (25 °C). SDS‐PAGE gels were used to determine abundance of the CO₂‐fixing enzyme, Rubisco. Leaf chlorophyll, nitrogen (N) and mass per unit leaf area (LMA) were also determined. For all species and T_growth, A_n at current atmospheric CO₂ partial pressure was Rubisco‐limited. Across all species, LMA decreased with increasing T_growth. Similarly, area‐based rates of V_cmax at a measurement T of 25 °C (V_cmax²⁵) linearly declined with increasing T_growth, linked to a concomitant decline in total leaf protein per unit leaf area and Rubisco as a percentage of leaf N. The decline in Rubisco constrained V_cmax and A_n for leaves developed at higher T_growth and resulted in poor predictions of photosynthesis by currently widely used models that do not account for T_growth‐mediated changes in Rubisco abundance that underpin the thermal acclimation response of photosynthesis in wet‐forest tree species. A new model is proposed that accounts for the effect of T_growth‐mediated declines in V_cmax²⁵ on A_n, complementing current photosynthetic thermal acclimation models that do not account for T sensitivity of V_cmax²⁵.     

Effects of logging on carbon dynamics of a jack pine forest in Saskatchewan,Canada

We calculated carbon budgets for a chronosequence of harvested jack pine (Pinus banksiana Lamb.) stands (0‐, 5‐, 10‐, and～29‐year‐old) and a～79‐year‐old stand that originated after wildfire. We measured total ecosystem C content (TEC), above‐, and belowground net primary productivity (NPP) for each stand. All values are reported in order for the 0‐, 5‐, 10‐, 29‐, and 79‐year‐old stands, respectively, for May 1999 through April 2000. Total annual NPP (NPP_T) for the stands (Mg C ha^?1 yr^?1±1 SD) was 0.9±0.3, 1.3±0.1, 2.7±0.6, 3.5±0.3, and 1.7±0.4. We correlated periodic soil surface CO₂ fluxes (R_S) with soil temperature to model annual R_S for the stands (Mg C ha^?1 yr^?1±1 SD) as 4.4±0.1, 2.4±0.0, 3.3±0.1, 5.7±0.3, and 3.2±0.2. We estimated net ecosystem productivity (NEP) as NPP_T minus R_H (where R_H was calculated using a Monte Carlo approach as coarse woody debris respiration plus 30–70% of total annual R_S). Excluding C losses during wood processing, NEP (Mg C ha^?1 yr^?1±1 SD) for the stands was estimated to be ?1.9±0.7, ?0.4±0.6, 0.4±0.9, 0.4±1.0, and ?0.2±0.7 (negative values indicate net sources to the atmosphere.) We also calculated NEP values from the changes in TEC among stands. Only the 0‐year‐old stand showed significantly different NEP between the two methods, suggesting a possible mismatch for the chronosequence. The spatial and methodological uncertainties allow us to say little for certain except that the stand becomes a source of C to the atmosphere following logging.     

Carbon sequestration in boreal jack pine stands following harvesting

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

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

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