Relative enhancement of photosynthesis and growth at elevated CO2 is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survivorship of dipterocarp seedlings in the deeply shaded understorey of South‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. To investigate the effect of elevated CO₂ upon photosynthesis and growth under sunflecks, seedlings of Shorealeprosula were grown in controlled environment conditions at ambient or elevated CO₂. Equal total daily photon flux density (PFD) (～7·7 mol m^?2 d^?1) was supplied as either uniform irradiance (～170 µmol m^?2 s^?1) or shade/fleck sequences (～30 µmol m^?2 s^?1/～525 µmol m^?2 s^?1). Photosynthesis and growth were enhanced by elevated CO₂ treatments but lower under flecked irradiance treatments. Acclimation of photosynthetic capacity occurred in response to elevated CO₂ but not flecked irradiance. Importantly, the relative enhancement effects of elevated CO₂ were greater under sunflecks (growth 60%, carbon gain 89%) compared with uniform irradiance (growth 25%, carbon gain 59%). This was driven by two factors: (1) greater efficiency of dynamic photosynthesis (photosynthetic induction gain and loss, post‐irradiance gas exchange); and (2) photosynthetic enhancement being greatest at very low PFD. This allowed improved carbon gain during both clusters of lightflecks (73%) and intervening periods of deep shade (99%). The relatively greater enhancement of growth and photosynthesis at elevated CO₂ under sunflecks has important potential consequences for seedling regeneration processes and hence forest structure and composition.     

The production of phytolith‐occluded carbon in China's forests: implications to biogeochemical carbon sequestration

The persistent terrestrial carbon sink regulates long‐term climate change, but its size, location, and mechanisms remain uncertain. One of the most promising terrestrial biogeochemical carbon sequestration mechanisms is the occlusion of carbon within phytoliths, the silicified features that deposit within plant tissues. Using phytolith content–biogenic silica content transfer function obtained from our investigation, in combination with published silica content and aboveground net primary productivity (ANPP) data of leaf litter and herb layer in China's forests, we estimated the production of phytolith‐occluded carbon (PhytOC) in China's forests. The present annual phytolith carbon sink in China's forests is 1.7 ± 0.4 Tg CO₂ yr ^{? 1}, 30% of which is contributed by bamboo because the production flux of PhytOC through tree leaf litter for bamboo is 3–80 times higher than that of other forest types. As a result of national and international bamboo afforestation and reforestation, the potential of phytolith carbon sink for China's forests and world's bamboo can reach 6.8 ± 1.5 and 27.0 ± 6.1 Tg CO₂ yr^?1, respectively. Forest management practices such as bamboo afforestation and reforestation may significantly enhance the long‐term terrestrial carbon sink and contribute to mitigation of global climate warming.     

Reconstruction and attribution of the carbon sink of European forests between 1950 and 2000

European forests are an important carbon sink; however, the relative contributions to this sink of climate, atmospheric CO₂ concentration ([CO₂]), nitrogen deposition and forest management are under debate. We attributed the European carbon sink in forests using ORCHIDEE‐FM, a process‐based vegetation model that differs from earlier versions of ORCHIDEE by its explicit representation of stand growth and idealized forest management. The model was applied on a grid across Europe to simulate changes in the net ecosystem productivity (NEP) of forests with and without changes in climate, [CO₂] and age structure, the three drivers represented in ORCHIDEE‐FM. The model simulates carbon stocks and volume increment that are comparable – root mean square error of 2 m³ ha^?1 yr^?1 and 1.7 kg C m^?2 respectively – with inventory‐derived estimates at country level for 20 European countries. Our simulations estimate a mean European forest NEP of 175 ± 52 g C m^?2 yr^?1 in the 1990s. The model simulation that is most consistent with inventory records provides an upwards trend of forest NEP of 1 ± 0.5 g C m^?2 yr^?2 between 1950 and 2000 across the EU 25. Furthermore, the method used for reconstructing past age structure was found to dominate its contribution to temporal trends in NEP. The potentially large fertilizing effect of nitrogen deposition cannot be told apart, as the model does not explicitly simulate the nitrogen cycle. Among the three drivers that were considered in this study, the fertilizing effect of increasing [CO₂] explains about 61% of the simulated trend, against 26% to changes in climate and 13% only to changes in forest age structure. The major role of [CO₂] at the continental scale is due to its homogeneous impact on net primary productivity (NPP). At the local scale, however, changes in climate and forest age structure often dominate trends in NEP by affecting NPP and heterotrophic respiration.     

Carbon balance indicator for forest bioenergy scenarios

A carbon (C) balance indicator is presented for the evaluation of forest bioenergy scenarios as a means to reduce greenhouse gas (GHG) emissions. A bioenergy‐intensive scenario with a greater harvest is compared to a baseline scenario. The relative carbon indicator (RC) is defined as the ratio between the difference in terrestrial C stocks – that is the C debt – and the difference in cumulative bioenergy harvest between the scenarios, over a selected time frame T. A value of zero indicates no C debt from additional biomass harvests, while a value of one indicates a C debt equal to the amount of additionally harvested biomass C. Multiplying the RC indicator by the smokestack emission factor of biomass (approximately 110 t CO₂/TJ) provides the net cumulative CO₂ emission factor of the biomass combustion as a function of T, allowing a direct comparison with the emission factors of comparable fossil fuels. The indicator is applied to bioenergy cases in Finland, where typically the rotation length of managed forests is long and the decay rate of harvest residues is slow. The country‐level examples illustrate that although Finnish forests remain as a C sink in each of the considered scenarios, the efforts of increasing forest bioenergy may still increase the atmospheric CO₂ concentrations in comparison with the baseline scenario and use of fossil fuels. The results also show that the net emission factor depends – besides on forest‐growth or residue‐decay dynamics – on the timing and evolution of harvests as well. Unlike for the constant fossil C emission factor, the temporal profile of bioenergy use is of great importance for the net emission factor of biomass.     

Partitioning direct and indirect human-induced effects on carbon sequestration of managed coniferous forests using model simulations and forest inventories

Temperate forest ecosystems have recently been identified as an important net sink in the global carbon budget. The factors responsible for the strength of the sinks and their permanence, however, are less evident. In this paper, we quantify the present carbon sequestration in Thuringian managed coniferous forests. We quantify the effects of indirect human‐induced environmental changes (increasing temperature, increasing atmospheric CO₂ concentration and nitrogen fertilization), during the last century using BIOME‐BGC, as well as the legacy effect of the current age‐class distribution (forest inventories and BIOME‐BGC). We focused on coniferous forests because these forests represent a large area of central European forests and detailed forest inventories were available. The model indicates that environmental changes induced an increase in biomass C accumulation for all age classes during the last 20 years (1982–2001). Young and old stands had the highest changes in the biomass C accumulation during this period. During the last century mature stands (older than 80 years) turned from being almost carbon neutral to carbon sinks. In high elevations nitrogen deposition explained most of the increase of net ecosystem production (NEP) of forests. CO₂ fertilization was the main factor increasing NEP of forests in the middle and low elevations. According to the model, at present, total biomass C accumulation in coniferous forests of Thuringia was estimated at 1.51 t C ha^?1 yr^?1 with an averaged annual NEP of 1.42 t C ha^?1 yr^?1 and total net biome production of 1.03 t C ha^?1 yr^?1 (accounting for harvest). The annual averaged biomass carbon balance (BCB: biomass accumulation rate‐harvest) was 1.12 t C ha^?1 yr^?1 (not including soil respiration), and was close to BCB from forest inventories (1.15 t C ha^?1 yr^?1). Indirect human impact resulted in 33% increase in modeled biomass carbon accumulation in coniferous forests in Thuringia during the last century. From the forest inventory data we estimated the legacy effect of the age‐class distribution to account for 17% of the inventory‐based sink. Isolating the environmental change effects showed that these effects can be large in a long‐term, managed conifer forest.     

Gypsy moth feeding in the canopy of a CO2-enriched mature forest

Rising atmospheric carbon dioxide (CO₂) concentration is expected to change plant tissue quality with important implications for plant–insect interactions. Taking advantage of canopy access by a crane and long‐term CO₂ enrichment (530 μ mol mol^?1) of a natural old‐growth forest (web‐free air carbon dioxide enrichment), we studied the responses of a generalist insect herbivore feeding in the canopy of tall trees. We found that relative growth rates (RGR) of gypsy moth (Lymantria dispar) were reduced by 30% in larvae fed on high CO₂‐exposed Quercus petraea, but increased by 29% when fed on high CO₂‐grown Carpinus betulus compared with control trees at ambient CO₂ (370 μ mol mol^?1). In Fagus sylvatica, there was a nonsignificant trend for reduced RGR under elevated CO₂. Tree species‐specific changes in starch to nitrogen ratio, water, and the concentrations of proteins, condensed and hydrolyzable tannins in response to elevated CO₂ were identified to correlate with altered RGR of gypsy moth larvae. Our data suggest that rising atmospheric CO₂ will have strong species‐specific effects on leaf chemical composition of canopy trees in natural forests leading to contrasting responses of herbivores such as those reported here. A future change in host tree preference seems likely with far‐ranging consequences for forest community dynamics.     

The effects of land use and climate change on the carbon cycle of Europe over the past 500 years

The long residence time of carbon in forests and soils means that both the current state and future behavior of the terrestrial biosphere are influenced by past variability in climate and anthropogenic land use. Over the last half‐millennium, European terrestrial ecosystems were affected by the cool temperatures of the Little Ice Age, rising CO₂ concentrations, and human induced deforestation and land abandonment. To quantify the importance of these processes, we performed a series of simulations with the LPJ dynamic vegetation model driven by reconstructed climate, land use, and CO₂ concentrations. Although land use change was the major control on the carbon inventory of Europe over the last 500 years, the current state of the terrestrial biosphere is largely controlled by land use change during the past century. Between 1500 and 2000, climate variability led to temporary sequestration events of up to 3 Pg, whereas increasing atmospheric CO₂ concentrations during the 20th century led to an increase in carbon storage of up to 15 Pg. Anthropogenic land use caused between 25 Pg of carbon emissions and 5 Pg of uptake over the same time period, depending on the historical and spatial pattern of past land use and the timing of the reversal from deforestation to afforestation during the last two centuries. None of the currently existing anthropogenic land use change datasets adequately capture the timing of the forest transition in most European countries as recorded in historical observations. Despite considerable uncertainty, our scenarios indicate that with limited management, extant European forests have the potential to absorb between 5 and 12 Pg of carbon at the present day.     

The contributions of land-use change, CO2 fertilization, and climate variability to the Eastern US carbon sink

Atmospheric measurements and land‐based inventories imply that terrestrial ecosystems in the northern hemisphere are taking up significant amounts of anthropogenic cabon dioxide (CO₂) emissions; however, there is considerable disagreement about the causes of this uptake, and its expected future trajectory. In this paper, we use the ecosystem demography (ED) model to quantify the contributions of disturbance history, CO₂ fertilization and climate variability to the past, current, and future terrestrial carbon fluxes in the Eastern United States. The simulations indicate that forest regrowth following agricultural abandonment accounts for uptake of 0.11 Pg C yr^?1 in the 1980s and 0.15 Pg C yr^?1 in the 1990s, and regrowth following forest harvesting accounts for an additional 0.1 Pg C yr^?1 of uptake during both these decades. The addition of CO₂ fertilization into the model simulations increases carbon uptake rates to 0.38 Pg C yr^?1 in the 1980s and 0.47 Pg C yr^?1 in the 1990s. Comparisons of predicted aboveground carbon uptake to regional‐scale forest inventory measurements indicate that the model's predictions in the absence of CO₂ fertilization are 14% lower than observed, while in the presence of CO₂ fertilization, predicted uptake rates are 28% larger than observed. Comparable results are obtained from comparisons of predicted total Net Ecosystem Productivity to the carbon fluxes observed at the Harvard Forest flux tower site and in model simulations free‐air CO₂ enrichment (FACE) experiments. These results imply that disturbance history is the principal mechanism responsible for current carbon uptake in the Eastern United States, and that conventional biogeochemical formulations of plant growth overestimate the response of plants to rising CO₂ levels. Model projections out to 2100 imply that the carbon uptake arising from forest regrowth will increasingly be dominated by forest regrowth following harvesting. Consequently, actual carbon storage declines to near zero by the end of the 21st century as the forest regrowth that has occurred since agricultural abandonment comes into equilibrium with the landscape's new disturbance regime. Incorporating interannual climate variability into the model simulations gives rise to large interannual variation in regional carbon fluxes, indicating that long‐term measurements are necessary to detect the signature of processes that give rise to long‐term uptake and storage.     

CO2 balance of boreal, temperate, and tropical forests derived from a global database

Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome‐specific carbon budgets; to re‐examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO₂ balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 °C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO₂ balance required the introduction of substantial biome‐specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non‐CO₂ carbon fluxes are not presently being adequately accounted for.     

How drought events during the last century have impacted biomass carbon in Amazonian rainforests

During the last two decades, inventory data show that droughts have reduced biomass carbon sink of the Amazon forest by causing mortality to exceed growth. However, process-based models have struggled to include drought-induced responses of growth and mortality and have not been evaluated against plot data. A process-based model, ORCHIDEE-CAN-NHA, including forest demography with tree cohorts, plant hydraulic architecture and drought-induced tree mortality, was applied over Amazonia rainforests forced by gridded climate fields and rising CO₂ from 1901 to 2019. The model reproduced the decelerating signal of net carbon sink and drought sensitivity of aboveground biomass (AGB) growth and mortality observed at forest plots across selected Amazon intact forests for 2005 and 2010. We predicted a larger mortality rate and a more negative sensitivity of the net carbon sink during the 2015/16 El Niño compared with the former droughts. 2015/16 was indeed the most severe drought since 1901 regarding both AGB loss and area experiencing a severe carbon loss. We found that even if climate change did increase mortality, elevated CO₂ contributed to balance the biomass mortality, since CO₂-induced stomatal closure reduces transpiration, thus, offsets increased transpiration from CO₂-induced higher foliage area.     

A multiyear synthesis of soil respiration responses to elevated atmospheric CO2 from four forest FACE experiments

The rapidly rising concentration of atmospheric CO₂ has the potential to alter forest and global carbon cycles by altering important processes that occur in soil. Forest soils contain the largest and longest lived carbon pools in terrestrial ecosystems and are therefore extremely important to the land–atmosphere exchange of carbon and future climate. Soil respiration is a sensitive integrator of many soil processes that control carbon storage in soil, and is therefore a good metric of changes to soil carbon cycling. Here, we summarize soil respiration data from four forest free‐air carbon dioxide enrichment (FACE) experiments in developing and established forests that have been exposed to elevated atmospheric [CO₂] (168 μL L^?1 average enrichment) for 2–6 years. The sites have similar experimental design and use similar methodology (closed‐path infrared gas analyzers) to measure soil respiration, but differ in species composition of the respective forest communities. We found that elevated atmospheric [CO₂] stimulated soil respiration at all sites, and this response persisted for up to 6 years. Young developing stands experienced greater stimulation than did more established stands, increasing 39% and 16%, respectively, averaged over all years and communities. Further, at sites that had more than one community, we found that species composition of the dominant trees was a major controller of the absolute soil CO₂ efflux and the degree of stimulation from CO₂ enrichment. Interestingly, we found that the temperature sensitivity of bulk soil respiration appeared to be unaffected by elevated atmospheric CO₂. These findings suggest that stage of stand development and species composition should be explicitly accounted for when extrapolating results from elevated CO₂ experiments or modeling forest and global carbon cycles.     

Interactive effects of nitrogen deposition, tropospheric ozone, elevated CO2 and land use history on the carbon dynamics of northern hardwood forests

Temperate forests are affected by a wide variety of environmental factors that stem from human industrial and agricultural activities. In the north‐eastern US, important change agents include tropospheric ozone, atmospheric nitrogen deposition, elevated CO₂, and historical human land use. Although each of these has received attention for its effects on forest carbon dynamics, integrated analyses that examine their combined effects are rare. To examine the relative importance of all of these factors on current forest growth and carbon balances, we included them individually and in combination in a forest ecosystem model that was applied over the period of 1700–2000 under different scenarios of air pollution and land use history. Results suggest that historical increases in CO₂ and N deposition have stimulated forest growth and carbon uptake, but to different degrees following agriculture and timber harvesting. These differences resulted from the effects of each land use scenario on soil C and N pools and on the resulting degree of growth limitations by carbon vs. nitrogen. Including tropospheric ozone in the simulations offset a substantial portion of the increases caused by CO₂ and N deposition. This result is particularly relevant given that ozone pollution is widespread across much of the world and because broad‐scale spatial patterns of ozone are coupled with patterns of nitrogen oxide emissions. This was demonstrated across the study region by a significant correlation between ozone exposure and rates of N deposition and suggests that the reduction of N‐induced carbon sinks by ozone may be a common phenomenon in other regions. Collectively, the combined effects of all physical and chemical factors we addressed produced growth estimates that were surprisingly similar to estimates obtained in the absence of any form of disturbance. The implication of this result is that intact forests may show relatively little evidence of altered growth since preindustrial times despite substantial changes in their physical and chemical environment.     

Risky future for Mediterranean forests unless they undergo extreme carbon fertilization

Forest performance is challenged by climate change but higher atmospheric [CO₂] (c_a) could help trees mitigate the negative effect of enhanced water stress. Forest projections using data assimilation with mechanistic models are a valuable tool to assess forest performance. Firstly, we used dendrochronological data from 12 Mediterranean tree species (six conifers and six broadleaves) to calibrate a process‐based vegetation model at 77 sites. Secondly, we conducted simulations of gross primary production (GPP) and radial growth using an ensemble of climate projections for the period 2010–2100 for the high‐emission RCP8.5 and low‐emission RCP2.6 scenarios. GPP and growth projections were simulated using climatic data from the two RCPs combined with (i) expected c_a; (ii) constant c_a = 390 ppm, to test a purely climate‐driven performance excluding compensation from carbon fertilization. The model accurately mimicked the growth trends since the 1950s when, despite increasing c_a, enhanced evaporative demands precluded a global net positive effect on growth. Modeled annual growth and GPP showed similar long‐term trends. Under RCP2.6 (i.e., temperatures below +2 °C with respect to preindustrial values), the forests showed resistance to future climate (as expressed by non‐negative trends in growth and GPP) except for some coniferous sites. Using exponentially growing c_a and climate as from RCP8.5, carbon fertilization overrode the negative effect of the highly constraining climatic conditions under that scenario. This effect was particularly evident above 500 ppm (which is already over +2 °C), which seems unrealistic and likely reflects model miss‐performance at high c_a above the calibration range. Thus, forest projections under RCP8.5 preventing carbon fertilization displayed very negative forest performance at the regional scale. This suggests that most of western Mediterranean forests would successfully acclimate to the coldest climate change scenario but be vulnerable to a climate warmer than +2 °C unless the trees developed an exaggerated fertilization response to [CO₂].     

Rising concentrations of atmospheric CO2 have increased growth in natural stands of quaking aspen (Populus tremuloides)

As atmospheric CO₂ levels rise, temperate and boreal forests in the Northern Hemisphere are gaining importance as carbon sinks. Quantification of that role, however, has been difficult due to the confounding effects of climate change. Recent large‐scale experiments with quaking aspen (Populus tremuloides), a dominant species in many northern forest ecosystems, indicate that elevated CO₂ levels can enhance net primary production. Field studies also reveal that droughts contribute to extensive aspen mortality. To complement this work, we analyzed how the growth of wild aspen clones in Wisconsin has responded to historical shifts in CO₂ and climate, accounting for age, genotype (microsatellite heterozygosity), and other factors. Aspen growth has increased an average of 53% over the past five decades, primarily in response to the 19.2% rise in ambient CO₂ levels. CO₂‐induced growth is particularly enhanced during periods of high moisture availability. The analysis accounts for the highly nonlinear changes in growth rate with age, and is unaffected by sex or location sampled. Growth also increases with individual heterozygosity, but this heterozygote advantage has not changed with rising levels of CO₂ or moisture. Thus, increases in future growth predicted from previous large‐scale, common‐garden work are already evident in this abundant and ecologically important tree species. Owing to aspen's role as a foundation species in many North American forest ecosystems, CO₂‐stimulated growth is likely to have repercussions for numerous associated species and ecosystem processes.     

Assessing forest growth across southwestern Oregon under a range of current and future global change scenarios using a process model, 3‐PG

With improvements in mapping regional distributions of vegetation using satellite‐derived information, there is an increasing interest in the assessment of current limitations on forest growth and in making projections of how productivity may be altered in response to changing climatic conditions and management policies. We utilised a simplified physiologically based process model (3‐PG) across a 54 000 km² mountainous region of southwestern Oregon, USA, to evaluate the degree to which maximum periodic mean annual increment (PAI) of forests could be predicted at a set of 448 forest inventory plots. The survey data were pooled into six broad forest types (coastal rain forest, interior coast range forest, mixed conifer, dry‐site Douglas‐fir, subalpine forest, and pine forest) and compared to the 3‐PG predictions at a spatial resolution of 1 km². We found good agreement (r² = 0.84) between mean PAI values of forest productivity for the six forest types with those obtained from field surveys. With confidence at this broader level of integration, we then ran model simulations to evaluate the constraints imposed by (i) soil fertility under current climatic conditions, (ii) the effect of doubling monthly precipitation across the region, and (iii) a widely used climatic change scenario that involves modifications in monthly mean temperatures and precipitation, as well as a doubling in atmospheric CO₂ concentrations. These analyses showed that optimum soil fertility would more than double growth, with the greatest response in the subalpine type and the least increase in the coastal rain forests. Doubling the precipitation increased productivity in the pine type (> 50%) with reduced responses elsewhere. The climate change scenario with doubled atmospheric CO₂ increased growth by 50% on average across all forest types, primarily as a result of a projected 33% increase in photosynthetic capacity. This modelling exercise indicates that, at a regional scale, a general relationship exists between simulated maximum leaf area index and maximum aboveground growth, supporting the contention that satellite‐derived estimates of leaf area index may be good measures of the potential productivity of temperate evergreen forests.     

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.     

Growth and phenology of mature temperate forest trees in elevated CO2

Are mature forest trees carbon limited at current CO₂ concentrations? Will ‘mid‐life’, 35 m tall deciduous trees grow faster in a CO₂‐enriched atmosphere? To answer these questions we exposed ca. 100‐year‐old temperate forest trees at the Swiss Canopy Crane site near Basel, Switzerland to a ca. 540 ppm CO₂ atmosphere using web‐FACE technology. Here, we report growth responses to elevated CO₂ for 11 tall trees (compared with 32 controls) of five species during the initial four treatment years. Tested across all trees, there was no CO₂ effect on stem basal area (BA) increment (neither when tested per year nor cumulatively for 4 years). In fact, the 4th year means were almost identical for the two groups. Stem growth data were standardized by pretreatment growth (5 years) in order to account for a priori individual differences in vigor. Although this experiment was not designed to test species specific effects, one species, the common European beech, Fagus sylvatica, showed a significant growth enhancement in the first year, which reoccurred during a centennial drought in the third year. None of the other dominant species (Quercus petraea, Carpinus betulus) showed a growth response to CO₂ in any of the 4 years or for all years together. The inclusion or exclusion of single individuals of Prunus avium and Tilia platyphyllos did not change the picture. In elevated CO₂, lateral branching in terminal shoots was higher in Fagus in 2002, when shoots developed from buds that were formed during the first season of CO₂ enrichment (2001), but there was no effect in later years and no change in lateral branching in any of the other species. In Quercus, there was a steady stimulation of leading shoot length in high‐CO₂ trees. Phenological variables (bud break, leaf fall, leaf duration) were highly species specific and were not affected by elevated CO₂ in any consistent way. Our 4‐year data set reflects a very dynamic and species‐specific response of tree growth to a step change in CO₂ supply. Stem growth after 4 years of exposure does not support the notion that mature forest trees will accrete wood biomass at faster rates in a future CO₂‐enriched atmosphere.     

Canopy-scale δ13C of photosynthetic and respiratory CO2 fluxes: observations in forest biomes across the United States

The δ¹³C values of atmospheric carbon dioxide (CO₂) can be used to partition global patterns of CO₂ source/sink relationships among terrestrial and oceanic ecosystems using the inversion technique. This approach is very sensitive to estimates of photosynthetic ¹³C discrimination by terrestrial vegetation (Δ_A), and depends on δ¹³C values of respired CO₂ fluxes (δ¹³C_R). Here we show that by combining two independent data streams – the stable isotope ratios of atmospheric CO₂ and eddy‐covariance CO₂ flux measurements – canopy scale estimates of Δ_A can be successfully derived in terrestrial ecosystems. We also present the first weekly dataset of seasonal variations in δ¹³C_R from dominant forest ecosystems in the United States between 2001 and 2003. Our observations indicate considerable summer‐time variation in the weekly value of δ¹³C_R within coniferous forests (4.0‰ and 5.4‰ at Wind River Canopy Crane Research Facility and Howland Forest, respectively, between May and September). The monthly mean values of δ¹³C_R showed a smaller range (2–3‰), which appeared to significantly correlate with soil water availability. Values of δ¹³C_R were less variable during the growing season at the deciduous forest (Harvard Forest). We suggest that the negative correlation between δ¹³C_R and soil moisture content observed in the two coniferous forests should represent a general ecosystem response to the changes in the distribution of water resources because of climate change. Shifts in δ¹³C_R and Δ_A could be of sufficient magnitude globally to impact partitioning calculations of CO₂ sinks between oceanic and terrestrial compartments.     

Old‐growth forests,carbon and climate change: Functions and management for tall open‐forests in two hotspots of temperate Australia

Abstract

The prognosis and utility under climate change are presented for two old‐growth, temperate forests in Australia, from ecological and carbon accounting perspectives. The tall open‐forests (TOFs) of south‐western Australia (SWA) are within Australia’s global biodiversity hotspot. The forest management and timber usage from the carbon‐dense old‐growth TOFs of Tasmania (TAS) have a high carbon efflux, rendering it a carbon hotspot. Under climate change the warmer, dryer climate in both areas will decrease carbon stocks directly; and indirectly through changes towards dryer forest types and through positive feedback. Near 2100, climate change will decrease soil organic carbon (SOC) significantly, e.g. by ～30% for SWA and at least 2% for TAS. The emissions from the next 20 years of logging old‐growth TOF in TAS, and conversion to harvesting cycles, will conservatively reach 66(±33) Mt‐CO₂‐equivalents in the long‐term – bolstering greenhouse gas emissions. Similar emissions will arise from rainforest SOC in TAS due to climate change. Careful management of old‐growth TOFs in these two hotspots, to help reduce carbon emissions and change in biodiversity, entails adopting approaches to forest, wood product and fire management which conserve old‐growth characteristics in forest stands. Plantation forestry on long‐cleared land and well‐targeted prescribed burning supplement effective carbon management.

Abbreviations: C, carbon; CBS, clearfell, burn and sow; CO₂‐e, CO₂ equivalents; CWD, coarse woody debris; DOC, dissolved organic carbon; GHG, greenhouse gas; Mt, megatonnes; SOC, soil organic carbon; SWA, south‐western Australia; SWAFR, Southwest Australian Floristic Region; TAS, Tasmania; TOF, tall open‐forest; t‐C ha^?1 yr^?1, tonnes of carbon per hectare per year     

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.