The carbon balance of tropical, temperate and boreal forests

Forest biomes are major reserves for terrestrial carbon, and major components of global primary productivity. The carbon balance of forests is determined by a number of component processes of carbon acquisition and carbon loss, and a small shift in the magnitude of these processes would have a large impact on the global carbon cycle. In this paper, we discuss the climatic influences on the carbon dynamics of boreal, temperate and tropical forests by presenting a new synthesis of micrometeorological, ecophysiological and forestry data, concentrating on three case-study sites. Historical changes in the carbon balance of each biome are also reviewed, and the evidence for a carbon sink in each forest biome and its likely behaviour under future global change are discussed. We conclude that there have been significant advances in determining the carbon balance of forests, but there are still critical uncertainties remaining, particularly in the behaviour of soil carbon stocks.     

The effect of climate anomalies and human ignition factor on wildfires in Russian boreal forests

Over the last few years anomalies in temperature and precipitation in northern Russia have been regarded as manifestations of climate change. During the same period exceptional forest fire seasons have been reported, prompting many authors to suggest that these in turn are due to climate change. In this paper, we examine the number and areal extent of forest fires across boreal Russia for the period 2002-2005 within two forest categories: 'intact forests' and 'non-intact forests'. Results show a far lower density of fire events in intact forests (5-14 times less) and that those events tend to be in the first 10 km buffer zone inside intact forest areas. Results also show that, during exceptional climatic years (2002 and 2003), fire event density is twice that found during normal years (2004 and 2005) and average areal extent of fire events (burned area) in intact forests is 2.5 times larger than normal. These results suggest that a majority of the fire events in boreal Russia are of human origin and a maximum of one-third of their impact (areal extension) can be attributed to climate anomalies alone, the rest being due to the combined effect of human disturbances and climate anomalies.     

Urban ecosystems and the North American carbon cycle

Approximately 75–80% of the population of North America currently lives in urban areas as defined by national census bureaus, and urbanization is continuing to increase. Future trajectories of fossil fuel emissions are associated with a high degree of uncertainty; however, if the activities of urban residents and the rate of urban land conversion can be captured in urban systems models, plausible emissions scenarios from major cities may be generated. Integrated land use and transportation models that simulate energy use and traffic‐related emissions are already in place in many North American cities. To these can be added a growing dataset of carbon gains and losses in vegetation and soils following urbanization, and a number of methods of validating urban carbon balance modeling, including top down atmospheric monitoring and urban ‘metabolic’ studies of whole ecosystem mass and energy flow. Here, we review the state of our understanding of urban areas as whole ecosystems with regard to carbon balance, including both drivers of fossil fuel emissions and carbon cycling in urban plants and soils. Interdisciplinary, whole‐ecosystem studies of the socioeconomic and biophysical factors that influence urban carbon cycles in a range of cities may greatly contribute to improving scenarios of future carbon balance at both continental and global scales.     

Vulnerability of carbon storage in North American boreal forests to wildfires during the 21st century

The boreal forest contains large reserves of carbon. Across this region, wildfires influence the temporal and spatial dynamics of carbon storage. In this study, we estimate fire emissions and changes in carbon storage for boreal North America over the 21st century. We use a gridded data set developed with a multivariate adaptive regression spline approach to determine how area burned varies each year with changing climatic and fuel moisture conditions. We apply the process‐based Terrestrial Ecosystem Model to evaluate the role of future fire on the carbon dynamics of boreal North America in the context of changing atmospheric carbon dioxide (CO₂) concentration and climate in the A2 and B2 emissions scenarios of the CGCM2 global climate model. Relative to the last decade of the 20th century, decadal total carbon emissions from fire increase by 2.5–4.4 times by 2091–2100, depending on the climate scenario and assumptions about CO₂ fertilization. Larger fire emissions occur with warmer climates or if CO₂ fertilization is assumed to occur. Despite the increases in fire emissions, our simulations indicate that boreal North America will be a carbon sink over the 21st century if CO₂ fertilization is assumed to occur in the future. In contrast, simulations excluding CO₂ fertilization over the same period indicate that the region will change to a carbon source to the atmosphere, with the source being 2.1 times greater under the warmer A2 scenario than the B2 scenario. To improve estimates of wildfire on terrestrial carbon dynamics in boreal North America, future studies should incorporate the role of dynamic vegetation to represent more accurately post‐fire successional processes, incorporate fire severity parameters that change in time and space, account for human influences through increased fire suppression, and integrate the role of other disturbances and their interactions with future fire regime.     

The organic carbon budget of a shallow arctic tundra lake on the Tuktoyaktuk Peninsula,N.W.T., Canada

The organic carbon cycle of a shallow, tundra lake (mean depth 1.45 m) was followed for 5 weeks of the open water period by examining CO₂ fluxes through benthic respiration and anaerobic decomposition, photosynthesis of benthic and phytoplankton communities and gas exchange at the air-water interface. Total photosynthesis (as consumption of carbon dioxide) was 37.5 mmole C m^–2 d^–1, 83% of which was benthic and macrophytic. By direct measurement benthic respiration exceeded benthic photosynthesis by 6.6 mmole C m^–2 d^–1. The lake lost 1.4 × 10⁶ moles C in two weeks after ice melted by degassing C0₂, and 6.8 mmole C m^–2 d^–1 (1.5 × 10⁶ moles) during the remainder of the open water period; 2.2 mmole C m² d^–1 of this was release Of CO₂ stored in the sediments by cryoconcentration the previous winter. Anaerobic microbial decomposition was only 4% of the benthic aerobic respiration rate of 38 mmole C m^–2 d^–1. An annual budget estimate for the lake indicated that 50% of the carbon was produced by the benthic community, 20% by phytoplankton, and 30% was allochthonous material. The relative contribution of allochthonous input was in accordance with measurement of the ¹⁵N of sedimented organic matter.     

The human carbon budget: an estimate of the spatial distribution of metabolic carbon consumption and release in the United States

Carbon dioxide is taken up by agricultural crops and released soon after during the consumption of agricultural commodities. The global net impact of this process on carbon flux to the atmosphere is negligible, but impact on the spatial distribution of carbon dioxide uptake and release across regions and continents is significant. To estimate the consumption and release of carbon by humans over the landscape, we developed a carbon budget for humans in the United States. The budget was derived from food commodity intake data for the US and from algorithms representing the metabolic processing of carbon by humans. Data on consumption, respiration, and waste of carbon by humans were distributed over the US using geospatial population data with a resolution of ~450 × 450 m. The average adult in the US contains about 21 kg C and consumes about 67 kg C year⁻¹ which is balanced by the annual release of about 59 kg C as expired CO₂, 7 kg C as feces and urine, and less than 1 kg C as flatus, sweat, and aromatic compounds. In 2000, an estimated 17.2 Tg C were consumed by the US population and 15.2 Tg C were expired to the atmosphere as CO₂. Historically, carbon stock in the US human population has increased between 1790 and 2006 from 0.06 Tg to 5.37 Tg. Displacement and release of total harvested carbon per capita in the US is nearly 12% of per capita fossil fuel emissions. Humans are using, storing, and transporting carbon about the Earth’s surface. Inclusion of these carbon dynamics in regional carbon budgets can improve our understanding of carbon sources and sinks.     

Density of organic carbon in soil at coniferous forest sites in southern Finland

More detailed knowledge of the density of organic carbon in soils of boreal forests is needed for accurate estimates of the size of this C stock. We investigated the effect of vegetation type and associated site fertility on the C density at 30 mature coniferous forest sites in southern Finland and evaluated the importance of deep layers to the total C store in the soil by extending the sampling at eight of the sites to the depth of ground water level (2.4–4.6 m). The C density in the organic horizon plus 1 m thick mineral soil layer ranged from 4.0 kg/m² to 11.9 kg/m², and, on the average, increased towards the more productive vegetation types. Between the depth of 1 m and the ground water level the C density averaged 1.3–2.4 kg/m² at the studied vegetation types and these layers represented 18–28% of the total stock of C in the soil. The results emphasize the importance of also considering these deep layers to correctly estimate the total amount of C in these soils. At the least fertile sites the soil contained about 30% more C than phytomass, whereas at the more fertile sites the amount of C in soil was about 10% less than the amount bound in vegetation.     

Overgrazing and soil carbon dynamics in eastern Inner Mongolia of China

Eastern Inner Mongolia of China is a typical ecotone between sandy forests and steppe. Little is known about the effect of overgrazing on carbon loss from soil in semiarid steppe and sandy forests of the north of China. The soil carbon parameters were measured in a 10,000 ha natural reserve in eastern Inner Mongolia of China (43°30′–43°36′N, 117°06′–117°16′E). Three situations were compared: primary protected (PP), moderately protected (MP) and highly degraded (HD). Soil and litter samples were recovered in spring and summer. Soil organic carbon (SOC) and CO₂–C values decreased from the PP (9.23 kg m⁻² and 157 g m⁻²) to the HD (1.69 kg m⁻² and 57 g m⁻²) sites whereas the C mineralization rate increased toward the less restored sites (1.06–2.37). Surface-litter C was different in both sites under protection (PP 648 and MP 408 g m⁻²), an was low at the HD site (17 g m⁻²). Leaves from woody species dominated the surface litter at the PP site, whereas grass material was predominant at the MP site. During summer, both CO₂–C and SOC decreased, whereas the C mineralization rate increased. We calculated that C loss since the introduction of cattle into the forest was 77 M g ha⁻¹, reaching a total of 1.1×10¹⁵ g for eastern Inner Mongolia. These values are higher than those caused by the conversion of steppe and other ecosystems into agriculture or cultivated pastures. The amount of C fixed at the PP site (650 g ha⁻¹year⁻¹) indicates that the sandy soils have a significant potential as atmospheric carbon sinks. Foundation item: The National Natural Science Foundation of China (No. 39900019, 30070129).     

Land use change and the global carbon cycle: the role of tropical soils

Millions of hectares of tropical forest are cleared annually for agriculture, pasture, shifting cultivation and timber. One result of these changes in land use is the release of CO₂ from the cleared vegetation and soils. Although there is uncertainty as to the size of this release, it appears to be a major source of atmospheric CO₂, second only to the release from the combustion of fossil fuels. This study estimates the release of CO₂ from tropical soils using a computer model that simulates land use change in the tropics and data on (1) the carbon content of forest soils before clearing; (2) the changes in the carbon content under the various types of land use; and (3) the area of forest converted to each use. It appears that the clearing and use of tropical soils affects their carbon content to a depth of about 40 cm. Soils of tropical closed forests contain approximately 6.7 kg C · m^-2; soils of tropical open forests contain approximately 5.2 kg C · m^-2 to this depth. The cultivation of tropical soils reduces their carbon content by 40% 5 yr after clearing; the use of these soils for pasture reduces it by about 20%. Logging in tropical forests appears to have little effect on soil carbon. The carbon content of soils used by shifting cultivators returns to the level found under primary forest about 35 yr after abandonment. The estimated net release of carbon from tropical soils due to land use change was 0.11–0.26 × 10¹⁵ g in 1980.     

Variable temperature sensitivity of soil organic carbon in North American forests

We investigated mean residence time (MRT) for soil organic carbon (SOC) sampled from paired hardwood and pine forests located along a 22 °C mean annual temperature (MAT) gradient in North America. We used acid hydrolysis fractionation, radiocarbon analyses, long-term laboratory incubations (525-d), and a three-pool model to describe the size and kinetics of the acid insoluble C (AIC), active and slow SOC fractions in soil. We found that active SOC was 2 ± 0.2% (mean ± SE) of total SOC, with an MRT of 33 ± 6 days that decreased strongly with increasing MAT. In contrast, MRT for slow SOC and AIC (70 ± 6% and 27 ± 6% of total SOC, respectively) ranged from decades to thousands of years, and neither was significantly related to MAT. The accumulation of AIC (as a percent of total SOC) was greater in hardwood than pine stands (36% and 21%, respectively) although the MRT for AIC was longer in pine stands. Based on these results, we suggest that the responsiveness of most SOC decomposition in upland forests to global warming will be less than currently modeled, but any shifts in vegetation from hardwood to pine may alter the size and MRT of SOC fractions.     

Earth Worm: Biomass Measurement: A Science Activity for Middle School

Ridgely, Robert S. A Guide to the Birds of Panama Princeton, N.J.: Princeton University Press, 1976 394 pp., $15.00. Reviewed by Karl L. Trever

Weiner, Michael A. Man's Useful Plants New York: Macmillan Co., 1976 137 pp., $6.95. Reviewed by Carolyn M. Levin

Bendick, Jeanne How Animals Behave New York: Parents Magazine Press, 1976 64 pp., $4.96. Reviewed by Robert Deitchman

Bayne, B. L., ed. Marine Mussels: The Ecology and Physiology New York: Cambridge University Press, 1976 506 pp., $49.50. Reviewed by Brother Nicholas Sullivan

Brown, Vinson The Explorer Naturalist Harrisburg, Pa.: Stackpole Books, 1976 288 pp., $10.00. Reviewed by Paul F. Connor

Sheldon, William G. The Wilderness Home of the Giant Panda Amherst: The University of Massachusetts Press, 1975 196 pp., $12.50. Reviewed by Harold W. Puffer

Rudolph, Joachim (translated by H. C. Grinter) Chemistry for the Modern Mind New York: Macmillan Publishing Co., 1973 359 pp., $8.95. Reviewed by Donald Schwartz     

Exceptional carbon uptake in European forests during the warm spring of 2007: a data–model analysis

Temperate and boreal forests undergo drastic functional changes in the springtime, shifting within a few weeks from net carbon (C) sources to net C sinks. Most of these changes are mediated by temperature. The autumn 2006–winter 2007 record warm period was followed by an exceptionally warm spring in Europe, making spring 2007 a good candidate for advances in the onset of the photosynthetically active period. An analysis of a decade of eddy covariance data from six European forests stands, which encompass a wide range of functional types (broadleaf evergreen, broadleaf deciduous, needleleaf evergreen) and a wide latitudinal band (from 44° to 62°N), revealed exceptional fluxes during spring 2007. Gross primary productivity (GPP) of spring 2007 was the maximum recorded in the decade examined for all sites but a Mediterranean evergreen forest (with a +40 to +130 gC m^?2 anomaly compared with the decadal mean over the January–May period). Total ecosystem respiration (TER) was also promoted during spring 2007, though less anomalous than GPP (with a +17 to +93 gC m^?2 anomaly over 5 months), leading to higher net uptake than the long‐term mean at all sites (+12 to +79 gC m^?2 anomaly over 5 months). A correlative analysis relating springtime C fluxes to simple phenological indices suggested spring C uptake and temperatures to be related. The CASTANEA process‐based model was used to disentangle the seasonality of climatic drivers (incoming radiation, air and soil temperatures) and biological drivers (canopy dynamics, thermal acclimation of photosynthesis to low temperatures) on spring C fluxes along the latitudinal gradient. A sensitivity analysis of model simulations evidenced the roles of (i) an exceptional early budburst combined with elevated air temperature in deciduous sites, and (ii) an early relief of winter thermal acclimation in coniferous sites for the promotion of 2007 spring assimilation.     

Modeling analysis of primary controls on net ecosystem productivity of seven boreal and temperate coniferous forests across a continental transect

Process‐based models are effective tools to synthesize and/or extrapolate measured carbon (C) exchanges from individual sites to large scales. In this study, we used a C‐ and nitrogen (N)‐cycle coupled ecosystem model named CN‐CLASS (Carbon Nitrogen‐Canadian Land Surface Scheme) to study the role of primary climatic controls and site‐specific C stocks on the net ecosystem productivity (NEP) of seven intermediate‐aged to mature coniferous forest sites across an east–west continental transect in Canada. The model was parameterized using a common set of parameters, except for two used in empirical canopy conductance–assimilation, and leaf area–sapwood relationships, and then validated using observed eddy covariance flux data. Leaf Rubisco‐N dynamics that are associated with soil–plant N cycling, and depend on canopy temperature, enabled the model to simulate site‐specific gross ecosystem productivity (GEP) reasonably well for all seven sites. Overall GEP simulations had relatively smaller differences compared with observations vs. ecosystem respiration (RE), which was the sum of many plant and soil components with larger variability and/or uncertainty associated with them. Both observed and simulated data showed that, on an annual basis, boreal forest sites were either carbon‐neutral or a weak C sink, ranging from 30 to 180 g C m^?2 yr^?1; while temperate forests were either a medium or strong C sink, ranging from 150 to 500 g C m^?2 yr^?1, depending on forest age and climatic regime. Model sensitivity tests illustrated that air temperature, among climate variables, and aboveground biomass, among major C stocks, were dominant factors impacting annual NEP. Vegetation biomass effects on annual GEP, RE and NEP showed similar patterns of variability at four boreal and three temperate forests. Air temperature showed different impacts on GEP and RE, and the response varied considerably from site to site. Higher solar radiation enhanced GEP, while precipitation differences had a minor effect. Magnitude of forest litter content and soil organic matter (SOM) affected RE. SOM also affected GEP, but only at low levels of SOM, because of low N mineralization that limited soil nutrient (N) availability. The results of this study will help to evaluate the impact of future climatic changes and/or forest C stock variations on C uptake and loss in forest ecosystems growing in diverse environments.     

Root production and mortality under elevated atmospheric carbon dioxide

An essential component of an understanding of carbon flux is the quantification of movement through the root carbon pool. Although estimates have been made using radiocarbon, the use of minirhizotrons provides a direct measurement of rates of root birth and death. We have measured root demographic parameters under a semi-natural grassland and for wheat. The grassland was studied along a natural altitudinal gradient in northern England, and similar turf from the site was grown in elevated CO₂ in solardomes. Root biomass was enhanced under elevated CO₂. Root birth and death rates were both increased to a similar extent in elevated CO₂, so that the throughput of carbon was greater than in ambient CO₂, but root half-lives were shorter under elevated CO₂ only under a Juncus/Nardus sward on a peaty gley soil, and not under a Festuca turf on a brown earth soil. In a separate experiment, wheat also responded to elevated CO₂ by increased root production, and there was a marked shift towards surface rooting: root development at a depth of 80–85 cm was both reduced and delayed. In conjunction with published results for trees, these data suggest that the impact of elevated CO₂ will be system-dependent, affecting the spatio-temporal pattern of root growth in some ecosystems and the rate of turnover in others. Turrnover is also sensitive to temperature, soil fertility and other environmental variables, all of which are likely to change in tandem with atmospheric CO₂ concentrations. Differences in turnover and time and location of rhizodeposition may have a large effect on rates of carbon cycling.     

The role of carbon dioxide in light-activated hydrogen production by Chlamydomonas reinhardtii

Light-activated hydrogen and oxygen evolution as a function of CO₂ concentration in helium were measured for the unicellular green alga Chlamydomonas reinhardtii. The concentrations were 58, 30, 0.8 and 0 ppm CO₂. The objective of these experiments was to study the differential affinity of CO₂/HCO₃^- for their respective Photosystem II and Calvin cycle binding sites vis-à-vis photoevolution of molecular oxygen and the competitive pathways of hydrogen photoevolution and CO₂ photoassimilation. The maximum rate of hydrogen evolution occurred at 0.8 ppm CO₂, whereas the maximum rate of oxygen evolution occurred at 58 ppm CO₂. The key result of this work is that the rate of photosynthetic hydrogen evolution can be increased by, at least partially, satisfying the Photosystem II CO₂/HCO₃^- binding site requirement without fully activating the Calvin-Benson CO₂ reduction pathway. Data are presented which plot the rates of hydrogen and oxygen evolution as functions of atmospheric CO₂ concentration in helium and light intensity. The stoichiometric ratio of hydrogen to oxygen changed from 0.1 at 58 ppm to approximately 2.5 at 0.8 ppm. A discussion of partitioning of photosynthetic reductant between the hydrogen/hydrogenase and Calvin-Benson cycle pathways is presented.Abbreviations PET photosynthetic electron transport - PS Photosystem     

The role of gap phase processes in the biomass dynamics of tropical forests

The responses of tropical forests to global anthropogenic disturbances remain poorly understood. Above-ground woody biomass in some tropical forest plots has increased over the past several decades, potentially reflecting a widespread response to increased resource availability, for example, due to elevated atmospheric CO2 and/or nutrient deposition. However, previous studies of biomass dynamics have not accounted for natural patterns of disturbance and gap phase regeneration, making it difficult to quantify the importance of environmental changes. Using spatially explicit census data from large (50 ha) inventory plots, we investigated the influence of gap phase processes on the biomass dynamics of four 'old-growth' tropical forests (Barro Colorado Island (BCI), Panama; Pasoh and Lambir, Malaysia; and Huai Kha Khaeng (HKK), Thailand). We show that biomass increases were gradual and concentrated in earlier-phase forest patches, while biomass losses were generally of greater magnitude but concentrated in rarer later-phase patches. We then estimate the rate of biomass change at each site independent of gap phase dynamics using reduced major axis regressions and ANCOVA tests. Above-ground woody biomass increased significantly at Pasoh (+0.72% yr(-1)) and decreased at HKK (-0.56% yr(-1)) independent of changes in gap phase but remained stable at both BCI and Lambir. We conclude that gap phase processes play an important role in the biomass dynamics of tropical forests, and that quantifying the role of gap phase processes will help improve our understanding of the factors driving changes in forest biomass as well as their place in the global carbon budget.     

A model of the long-term response of carbon allocation and productivity of forests to increased CO2concentration and nitrogen deposition

We present a simple theoretical analysis of the long term response of forest growth and carbon allocation to increased atmospheric [CO₂] and N deposition. Our analysis is based on a recent model which predicts that plant light-use efficiency increases with [CO₂] but is independent of plant N supply. We combine that model with simple assumptions for nitrogen fluxes in the soil. A quasi-equilibrium analysis of the short term tree and soil pools is then used to develop a simple graphical depiction of the long term carbon and nitrogen supply constraints on total growth, stem growth and foliar allocation. Our results suggest that long-term growth responses to [CO₂] and N deposition depend strongly on the extent to which stem allocation and foliage allocation are coupled. At one extreme (‘no coupling’), when stem allocation is fixed and independent of foliage allocation, there is no response of total growth or stem growth to increased [CO₂] unless N deposition increases. At the other extreme (‘linear coupling’), when stem allocation is proportional to foliage allocation, there is a significant long-term increase in total growth following a doubling of [CO₂], even when N deposition is unchanged, but stem growth decreases because of a long-term decrease in foliage allocation. For both types of coupling, total growth and stem growth increase with increasing N deposition. In the case of linear coupling, however, the N deposition response of stem growth is significantly larger than that of total growth, because of a long-term increase in foliage allocation. We compare our results with those obtained previously from an alternative model of canopy light-use efficiency involving a dependence on the foliar N:C ratio in addition to [CO₂]. Our results highlight the need for more experimental information on (i) the extent to which canopy light-use efficiency is independent of N supply, and (ii) the relationship between foliage allocation and stem allocation.     

Methanogenic activity of woody plants

Methane production in trunks of living and dead trees was demonstrated. Forest trees are one of sources for this gas emission into the atmosphere. Quantitative evaluation of the methagenic activity of living wood and that digested by xylotrophic fungi is presented.     

Belowground carbon storage of Micronesian mangrove forests

Belowground carbon storage was examined for mangrove forests on Pohnpei Island, Micronesia. Stored carbon in a coral reef-type mangrove habitat consisting of a 2 m thick mangrove peat layer, which is a type of mangrove habitat in tropical Pacific islands, was estimated at 1300 t C ha^–1. The carbon burial rate during the phase of gradual sea-level rise, which was calculated at 93 g m^–2 year^–1 between 1800 and 1380 years bp using the medians of the radiocarbon ages, was significantly higher than that between 1380 years bp and present in a stable sea-level phase.