Testing for a CO2 fertilization effect on growth of Canadian boreal forests

The CO2 fertilization hypothesis stipulates that rising atmospheric CO2 has a direct positive effect on net primary productivity (NPP), with experimental evidence suggesting a 23% growth enhancement with a doubling of CO2. Here, we test this hypothesis by comparing a bioclimatic model simulation of NPP over the twentieth century against tree growth increment (TGI) data of 192 Pinus banksiana trees from the Duck Mountain Provincial Forest in Manitoba, Canada. We postulate that, if a CO2 fertilization effect has occurred, climatically driven simulations of NPP and TGI will diverge with increasing CO2. We use a two-level scaling approach to simulate NPP. A leaf-level model is first used to simulate high-frequency responses to climate variability. A canopy-level model of NPP is then adjusted to the aggregated leaf-level results and used to simulate yearly plot-level NPP. Neither model accounts for CO2 fertilization. The climatically driven simulations of NPP for 1912–2000 are effective for tracking the measured year-to-year variations in TGI, with 47.2% of the variance in TGI reproduced by the simulation. In addition, the simulation reproduces without divergence the positive linear trend detected in TGI over the same period. Our results therefore do not support the attribution of a portion of the historical linear trend in TGI to CO2 fertilization at the level suggested by current experimental evidence. A sensitivity analysis done by adding an expected CO2 fertilization effect to simulations suggests that the detection limit of the study is for a 14% growth increment with a doubling of atmospheric CO2 concentration.

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title: Response of Forest Trees to Increased Atmospheric CO 2
abstract: The CO 2 fertilization hypothesis stipulates that rising atmospheric CO 2 has a positive effect on tree growth due to increasing availability of carbon. The objective of this paper is to compare the recent literature related to both field CO 2 -enriched experiments with trees and empirical dendrochronological studies detecting CO 2 fertilization effects in tree-rings. This will allow evaluation of tree growth responses to atmospheric CO 2 enrichment by combining evidence from both ecophysiology and tree-ring research. Based on considerable experimental evidence of direct CO 2 fertilization effect (increased photosynthesis, water use efficiency, and above- and belowground biomass), and predications from the interactions of enriched CO 2 with temperature, nitrogen and drought, we propose that warm, moderately drought-stressed ecosystems with an ample nitrogen supply might be the most CO 2 responsive ecosystems. Empirical tree-ring studies took the following three viewpoints on detecting CO 2 fertilization effect in tree-rings: 1) finding evidence of CO 2 fertilization effect in tree-rings, 2) attributing growth enhancement to favorable climate rather than atmospheric CO 2 enrichment, and 3) considering that tree growth enhancement might be caused by synergistic effects of several factors such as favorable climate change, CO 2 fertilization, and anthropogenic atmospheric deposition (e.g., nitrogen). At temperature-limiting sites such as high elevations, nonfindings of CO 2 fertilization evidence could be ascribed to the following possibilities: 1) cold temperatures, a short season of cambial division, and nitrogen deficiency that preclude a direct CO 2 response, 2) old trees past half of their maximum life expectancy and consequently only a small increase in biomass increment due to CO 2 fertilization effect might be diminished, 3) the elimination of age/size-related trends by statistical detrending of tree-ring series that might remove some long-term CO 2 -related trends in tree-rings, and 4) carbon partitioning and growth within a plant that is species-specific. Our review supports the atmospheric CO 2 fertilization effect hypothesis, at least in trees growing in semi-arid or arid conditions because the drought-stressed trees could benefit from increased water use efficiency to enhance growth.

title: What have we learned from 15 years of free-air CO 2 enrichment ( FACE ) ? A meta-analytic review of the responses of photosynthesis , canopy properties and plant production to rising CO 2
abstract: Free-air CO 2 enrichment (FACE) experiments allow study of the effects of elevated [CO 2 ] on plants and ecosystems grown under natural conditions without enclosure. Data from 120 primary, peer-reviewed articles describing physiology and production in the 12 large-scale FACE experiments (475–600 ppm) were collected and summarized using meta-analytic techniques. The results confirm some results from previous chamber experiments: light-saturated carbon uptake, diurnal C assimilation, growth and above-ground production increased, while specific leaf area and stomatal conductance decreased in elevated [CO 2 ]. There were differences in FACE. Trees were more responsive than herbaceous species to elevated [CO 2 ]. Grain crop yields increased far less than anticipated from prior enclosure studies. The broad direction of change in photosynthesis and production in elevated [CO 2 ] may be similar in FACE and enclosure studies, but there are major quantitative differences: trees were more responsive than other functional types; C 4 species showed little response; and the reduction in plant nitrogen was small and largely accounted for by decreased Rubisco. The results from this review may provide the most plausible estimates of Author for correspondence: Stephen P. Long Tel: + 1 217 333 2487

title: Ecosystem effects of CO 2 concentration: evidence from past climates
abstract: Atmospheric CO2 concentration has varied from minima of 170–200 ppm in glacials to maxima of 280–300 ppm in the recent interglacials. Photosynthesis by C3 plants is highly sensitive to CO2 concentration variations in this range. Physiological consequences of the CO2 changes should therefore be discernible in palaeodata. Several lines of evidence support this expectation. Reduced terrestrial carbon storage during glacials, indicated by the shift in stable isotope composition of dissolved inorganic carbon in the ocean, cannot be explained by climate or sea-level changes. It is however consistent with predictions of current process-based models that propagate known physiological CO2 effects into net primary production at the ecosystem scale. Restricted forest cover during glacial periods, indicated by pollen assemblages dominated by non-arboreal taxa, cannot be reproduced accurately by palaeoclimate models unless CO2 effects on C3-C4 plant competition are also modelled. It follows that methods to reconstruct climate from palaeodata should account for CO2 concentration changes. When they do so, they yield results more consistent with palaeoclimate models. In conclusion, the palaeorecord of the Late Quaternary, interpreted with the help of climate and ecosystem models, provides evidence that CO2 effects at the ecosystem scale are neither trivial nor transient.

title: Forest response to elevated CO2 is conserved across a broad range of productivity.
abstract: Climate change predictions derived from coupled carbon-climate models are highly dependent on assumptions about feedbacks between the biosphere and atmosphere. One critical feedback occurs if C uptake by the biosphere increases in response to the fossil-fuel driven increase in atmospheric [CO(2)] ("CO(2) fertilization"), thereby slowing the rate of increase in atmospheric [CO(2)]. Carbon exchanges between the terrestrial biosphere and atmosphere are often first represented in models as net primary productivity (NPP). However, the contribution of CO(2) fertilization to the future global C cycle has been uncertain, especially in forest ecosystems that dominate global NPP, and models that include a feedback between terrestrial biosphere metabolism and atmospheric [CO(2)] are poorly constrained by experimental evidence. We analyzed the response of NPP to elevated CO(2) ( approximately 550 ppm) in four free-air CO(2) enrichment experiments in forest stands. We show that the response of forest NPP to elevated [CO(2)] is highly conserved across a broad range of productivity, with a stimulation at the median of 23 +/- 2%. At low leaf area indices, a large portion of the response was attributable to increased light absorption, but as leaf area indices increased, the response to elevated [CO(2)] was wholly caused by increased light-use efficiency. The surprising consistency of response across diverse sites provides a benchmark to evaluate predictions of ecosystem and global models and allows us now to focus on unresolved questions about carbon partitioning and retention, and spatial variation in NPP response caused by availability of other growth limiting resources.

title: Simulated responses of Pinus halepensis forest productivity to climatic change and CO2 increase using a statistical model
abstract: Tree ring chronologies provide long-term records of growth in natural environmental conditions and may be used to evaluate impacts of climatic change and CO2 increase on forest productivity. This study focuses on 21 Pinus halepensis forest stands in calcareous Provence (in the south-east of France). A chronology of net primary productivity (NPP) both for the 20th century and for each stand was estimated using tree ring data (width and density). The response of each stand to climate in terms of NPP was statistically modelled using response functions. Anomalies between estimated NPP and NPP reconstructed by response functions were calculated to evaluate the fertilising effect of CO2 increase on tree growth. The changes in anomalies during the 20th century were attributed to the effect of CO2 increase. A multiplying factor (beta) linking CO2 concentration and stand productivity was then calculated, on the basis of the trend observed during the 20th century. In this study, the value of the beta factor obtained under natural conditions (beta = 0.50) is consistent with those from controlled CO2 enrichment experiments. Both response functions and the beta factor were used to predict NPP changes for a 2 X CO2 scenario. The 2 X CO2 climate was obtained using predictions from Meteo France's ARPEGE atmospheric general circulation model (AGCM) downscaled to Marseilles meteorological station. NPP increased significantly for nine stands solely when the climatic effect was taken into account. The main factors responsible for this enhancement were increased winter and early spring temperatures. When the fertilising effect of the CO2 increase was added, NPP was significantly enhanced for 14 stands (i.e. NPP enhancement ranged from 8% to 55%). Although the effects of global change were slightly detectable during the 20th century, their acceleration is likely to lead to great changes in the future productivity of P. halepensis forests.

title: Using a biogeochemistry model in simulating forests productivity responses to climatic change and [CO2] increase: example of Pinus halepensis in Provence (south-east France)
abstract: Tree-ring chronologies provide long-term records of growth in natural environmental conditions and may be used to evaluate the impacts of climatic change and [CO2] increase on forest productivity. This study focuses on 21 Pinus halepensis forest stands in calcareous Provence (in the south-east France). Changes in productivity are simulated using the global biogeochemistry model BIOME3, that we have adapted to run with chronological data. Tree-ring data (width and density) were used to estimate, for each stand, an observed series of changes in productivity. Simulated and observed productivity changes are then compared to validate the chronological biogeochemistry model BIOME3C. Variations in productivity were well reconstructed at 15 sites. After this validation, BIOME3C was used to simulate forest productivity changes for a 2 x CO2 scenario. The 2 x CO2 climate used as input was obtained using results from Meteo-France's ARPEGE atmospheric general circulation model (ALCM), downscaled to local meteorological stations. Productivity increases moderately for all stands (from 17 to 24%) when climatic changes alone were taken into account. The main factor responsible for this increase is a reduction in summer drought severity. Productivity increases highly for all stands (from 72 to 86%) when the physiological fertilising effect of the [CO2] increase is considered separately. When both climatic changes and the [CO2] increase were taken into account, productivity increases highly, from 107% (for Moustier) to 141 % (for La Ciotat). The direct fertilising effect of [CO2] increase has a greater influence on the forest stands productivity than the indirect climatic changes effect. These results also exhibit the importance of the synergy between the effects of climate change and [CO2] increase, as the increase in productivity resulting from the combined effects are more than the sum of the individual CO2 and climate effects. Although the detected effects of global change during the 20th century were slight, acceleration of these changes is likely to lead to great changes in the future productivity of P. halepensis forests.

title: Annual carbon balance of Canada's forests during 1895-1996.
abstract: This paper reports annual carbon (C) balance of Canada's forests during 1895–1996 estimated using the Integrated Terrestrial Ecosystem C-budget model (InTEC) [Chen et al. this issue]. During 1895–1910, Canada's forests were small sources of 30±15 Tg C yr−1 due to large disturbances (forest fire, insect-induced mortality, and harvest) in late nineteenth century. The forests became large sinks of 170±85 Tg C yr−1 during 1930–1970, owing to forest regrowth in previously disturbed areas and growth stimulation by nondisturbance factors such as climate, atmospheric CO2 concentration, and N deposition. In recent decades (1980–1996), Canada's forests have been moderate sinks of 50±25 Tg C yr−1, as a result of a tradeoff between the negative effects of increased disturbances and positive effects of nondisturbance factors. The nondisturbance factors, in order of importance, are (1) atmospheric N deposition (measured by a national monitoring network), (2) net N mineralization and fixation (estimated from temperature and precipitation records), (3) growing season length increase (estimated from spring air temperature records), and (4) CO2 fertilization (estimated from CO2 records using a leaf-level photosynthesis model). The magnitudes of modeled nondisturbance effects are consistent with simulation results by the Carnegie-Ames-Stanford Approach (CASA) and are also in broad agreement with flux measurements above mature forest stands at several locations in Canada. Results for the disturbance effects agree with a previous study [Kurz and Apps, 1996]. The overall C balance from InTEC generally agrees with that derived from tree ring data [Auclair and Bedford, 1997] and from forest inventories. The combination of our result and that of Houghton et al. [1999] for the United States suggests that North America (> 15°N) was probably a C sink of 0.2-0.5 Pg C yr±−1 during 1980s, much less than that of 1.7 Pg C yr±−1 estimated by Fan et al. [1998] using an atmospheric inversion method.

title: Simulating Picea schrenkiana forest productivity under climatic changes and atmospheric CO2 increase in Tianshan Mountains, Xinjiang Autonomous Region, China
abstract: A process-based model, BIOME-BGC, was used to investigate the response of Picea schrenkiana forest to future climate changes and atmospheric carbon dioxide (CO2) concentration increases in the Tianshan Mountains of northwestern China. The model was validated by comparing simulated net primary productivity (NPP) under current climatic conditions with independent field-measured data. Then the model was used to predict P. schrenkiana forest productivity response to the different climatic and CO2 change scenarios. The results showed that NPP increased moderately (about 18.6%) for all sites when both temperature and precipitation changes were taken into account. The main factor contributing to NPP increase was an increase in precipitation, which tended to alleviate moisture stress for P. schrenkiana forest growth. NPP increase was relatively low (about 2.7%) when the physiological fertilizing effect of the doubled atmospheric CO2 concentration was considered alone, in part because of nitrogen limitation and low temperatures. When both climatic changes and doubling of atmospheric CO2 concentration were taken into account, NPP increased more dramatically (from 26.4% to 37.2%). The results also indicated that the interactive effects of climate and CO2 changes were not a simple additive combination of the individual responses. The results provided support for theview that forest response to climatic change depends on local site conditions.

title: Quantifying the response of forest carbon balance to future climate change in Northeastern China: Model validation and prediction
abstract: In this study, we report on the validation of process-based forest growth and carbon and nitrogen model of TRIPLEX against observed data, and the use of the model to investigate the potential impacts and interaction of climate change and increasing atmospheric CO2 on forest net primary productivity (NPP) and carbon budgets in northeast of China. The model validation results show that the simulated tree total volume, NPP, total biomass and soil carbon are consistent with observed data across the Northeast of China, demonstrating that the improved TRIPLEX model is able to simulate forest growth and carbon dynamics of the boreal and temperate forest ecosystems at regional scale. The climate change would increase forest NPP and biomass carbon but decrease overall soil carbon under all three climate change scenarios. The combined effects of climate change and CO2 fertilization on the increase of NPP were estimated to be 10–12% for 2030s and 28–37% in 2090s. The simulated effects of CO2 fertilization significantly offset the soil carbon loss due to climate change alone. Overall, future climate change and increasing atmospheric CO2 would have a significant impact on the forest ecosystems of Northeastern China.

title: Global Vegetation and Climate Change due to Future Increases in CO2 as Projected by a Fully Coupled Model with Dynamic Vegetation
abstract: Transient simulations are presented of future climate and vegetation associated with continued rising levels of CO2. The model is a fully coupled atmosphere–ocean–land–ice model with dynamic vegetation. The impacts of the radiative and physiological forcing of CO2 are diagnosed, along with the role of vegetation feedbacks. While the radiative effect of rising CO2 produces most of the warming, the physiological effect contributes additional warming by weakening the hydrologic cycle through reduced evapotranspiration. Both effects cause drying over tropical rain forests, while the radiative effect enhances Arctic and Indonesian precipitation. A global greening trend is simulated primarily due to the physiological effect, with an increase in photosynthesis and total tree cover associated with enhanced water-use efficiency. In particular, tree cover is enhanced by the physiological effect over moisture-limited regions. Over Amazonia, South Africa, and Australia, the radiative forcing produces soil dr...

title: Carbon Flux and Growth in Mature Deciduous Forest Trees Exposed to Elevated CO2
abstract: Whether rising atmospheric carbon dioxide (CO2) concentrations will cause forests to grow faster and store more carbon is an open question. Using free air CO2 release in combination with a canopy crane, we found an immediate and sustained enhancement of carbon flux through 35-meter-tall temperate forest trees when exposed to elevated CO2. However, there was no overall stimulation in stem growth and leaf litter production after 4 years. Photosynthetic capacity was not reduced, leaf chemistry changes were minor, and tree species differed in their responses. Although growing vigorously, these trees did not accrete more biomass carbon in stems in response to elevated CO2, thus challenging projections of growth responses derived from tests with smaller trees.

title: Contributions of land-use history to carbon accumulation in U.S. forests.
abstract: Carbon accumulation in forests has been attributed to historical changes in land use and the enhancement of tree growth by CO2 fertilization, N deposition, and climate change. The relative contribution of land use and growth enhancement is estimated by using inventory data from five states spanning a latitudinal gradient in the eastern United States. Land use is the dominant factor governing the rate of carbon accumulation in these states, with growth enhancement contributing far less than previously reported. The estimated fraction of aboveground net ecosystem production due to growth enhancement is 2.0 +/- 4.4%, with the remainder due to land use.

title: Nitrogen Limitation is Reducing the Enhancement of NPP by Elevated CO2 in a Deciduous Forest
abstract: Accurate model representation of the long-term response of forested ecosystems to elevated atmospheric CO2 concentrations (eCO2) is important for predictions of future concentrations of CO2. For biogeochemical models that predict the response of net primary productivity (NPP) to eCO2, free-air CO2 enrichment (FACE) experiments provide the only source of data for comparison. A synthesis of forest FACE experiments reported a 23% increase in NPP in eCO2, and this result has been used as a model benchmark. Here, we provide new evidence from a FACE experiment in a deciduous forest in Tennessee that N limitation has significantly reduced the stimulation of NPP by eCO2, consistent with predictions from ecosystem and global models that incorporate N feedbacks. The Liquidambar styraciflua stand has been exposed to current ambient atmospheric CO2 or air enriched with CO2 to 550 ppm since 1998. Results from the first 6 years of the experiment indicated that NPP was significantly enhanced by eCO2 and that this was a consistent and sustained response. Now, with 10 years of data, our analysis must be revised. The response of NPP to eCO2 has declined from 24% in 2001-2003 to 9% in 2007. The diminishing response to eCO2 since 2004 coincides with decliningmore » NPP in ambient CO2 plots. Productivity of this forest stand is limited by N availability, and the steady decline in forest NPP is closely related to changes in the N economy, as evidenced by declining foliar N concentrations. There is a strong linear relationship between foliar [N] and NPP, and the steeper slope in eCO2 indicates that the NPP response to eCO2 should diminish as foliar N declines. Increased fine-root production and root proliferation deeper in the soil have sustained N uptake, but not to an extent sufficient to benefit aboveground production. The mechanistic basis of the N effect on NPP resides in the photosynthetic machinery. The linear relationships between Jmax and Vcmax with foliar [N] did not change from 1998 to 2008 or in response to eCO2; hence, lower foliar [N] resulted in significant reductions in Jmax, Vcmax, and photosynthesis over time and in eCO2. It is not yet clear whether foliar [N] and NPP will continue to decline or have reached a new steady state indicative of long-term forest response to eCO2. These results are consistent with ecosystem models, which suggest that the NPP response to eCO2 will include a transient increase in NPP followed by a decline to a lower level when fast C and N pools reach quasi-equilibrium at eCO2. Our results also are consistent with optimization models of carbon-water-nitrogen economy, which suggest eCO2 leads to increased fine-root production, declining foliar [N], and diminishing enhancement of aboveground production. At a larger scale, a model incorporating N feedbacks (CLM3-CN) predicts a much smaller enhancement of NPP in eCO2 than the CLM3-CASA model without such a feedback. When applied across the terrestrial biosphere, the smaller CO2 fertilization effect has important implications for the pace of climate change.« less

title: CO2 Enhancement of Forest Productivity Constrained by Limited Nitrogen Availability
abstract: Stimulation of terrestrial plant production by rising CO2 concentration is projected to reduce the airborne fraction of anthropogenic CO2 emissions. Coupled climate–carbon cycle models are sensitive to this negative feedback on atmospheric CO2, but model projections are uncertain because of the expectation that feedbacks through the nitrogen (N) cycle will reduce this so-called CO2 fertilization effect. We assessed whether N limitation caused a reduced stimulation of net primary productivity (NPP) by elevated atmospheric CO2 concentration over 11 y in a free-air CO2 enrichment (FACE) experiment in a deciduous Liquidambar styraciflua (sweetgum) forest stand in Tennessee. During the first 6 y of the experiment, NPP was significantly enhanced in forest plots exposed to 550 ppm CO2 compared with NPP in plots in current ambient CO2, and this was a consistent and sustained response. However, the enhancement of NPP under elevated CO2 declined from 24% in 2001–2003 to 9% in 2008. Global analyses that assume a sustained CO2 fertilization effect are no longer supported by this FACE experiment. N budget analysis supports the premise that N availability was limiting to tree growth and declining over time —an expected consequence of stand development, which was exacerbated by elevated CO2. Leafand stand-level observations provide mechanistic evidence that declining N availability constrained the tree response to elevated CO2; these observations are consistent with stand-level model projections. This FACE experiment provides strong rationale and process understanding for incorporating N limitation and N feedback effects in ecosystem and global models used in climate change assessments.

title: Soil fertility limits carbon sequestration by forest ecosystems in a CO2-enriched atmosphere
abstract: Northern mid-latitude forests are a large terrestrial carbon sink. Ignoring nutrient limitations, large increases in carbon sequestration from carbon dioxide (CO2) fertilization are expected in these forests. Yet, forests are usually relegated to sites of moderate to poor fertility, where tree growth is often limited by nutrient supply, in particular nitrogen. Here we present evidence that estimates of increases in carbon sequestration of forests, which is expected to partially compensate for increasing CO2 in the atmosphere, are unduly optimistic. In two forest experiments on maturing pines exposed to elevated atmospheric CO2, the CO2-induced biomass carbon increment without added nutrients was undetectable at a nutritionally poor site, and the stimulation at a nutritionally moderate site was transient, stabilizing at a marginal gain after three years. However, a large synergistic gain from higher CO2 and nutrients was detected with nutrients added. This gain was even larger at the poor site (threefold higher than the expected additive effect) than at the moderate site (twofold higher). Thus, fertility can restrain the response of wood carbon sequestration to increased atmospheric CO2. Assessment of future carbon sequestration should consider the limitations imposed by soil fertility, as well as interactions with nitrogen deposition.

title: Simulating the effects of climatic variation on stem carbon accumulation of a ponderosa pine stand: comparison with annual growth increment data.
abstract: Simulation models of ecosystem processes may be necessary to separate the long-term effects of climate change on forest productivity from the effects of year-to-year variations in climate. The objective of this study was to compare simulated annual stem growth with measured annual stem growth from 1930 to 1982 for a uniform stand of ponderosa pine (Pinus ponderosa Dougl.) in Montana, USA. The model, FOREST-BGC, was used to simulate growth assuming leaf area index (LAI) was either constant or increasing. The measured stem annual growth increased exponentially over time; the differences between the simulated and measured stem carbon accumulations were not large. Growth trends were removed from both the measured and simulated annual increments of stem carbon to enhance the year-to-year variations in growth resulting from climate. The detrended increments from the increasing LAI simulation fit the detrended increments of the stand data over time with an R(2) of 0.47; the R(2) increased to 0.65 when the previous year's simulated detrended increment was included with the current year's simulated increment to account for autocorrelation. Stepwise multiple linear regression of the detrended increments of the stand data versus monthly meteorological variables had an R(2) of 0.37, and the R(2) increased to 0.47 when the previous year's meteorological data were included to account for autocorrelation. Thus, FOREST-BGC was more sensitive to the effects of year-to-year climate variation on annual stem growth than were multiple linear regression models.

title: A generalised model of forest productivity using simplified concepts of radiation-use efficiency, carbon balance and partitioning
abstract: This paper describes a stand growth model, based on physiological processes, which incorporates a number of steps and procedures that have allowed considerable simplification relative to extant process-based models. The model, called 3-PG (use of Physiological Principles in Predicting Growth), calculates total carbon fixed (gross primary production; PG) from utilizable, absorbed photosynthetically active radiation (φp.a.u.), obtained by correcting the photosynthetically active radiation absorbed by the forest canopy (φp.a.) for the effects of soil drought, atmospheric vapour pressure deficits and stand age. PG is obtained from φp.a.u. and the canopy quantum efficiency, values of which are becoming available. The ratio of net (PN) to gross primary production is emerging as relatively constant for trees. This eliminates the need to calculate respiration and is used to estimate PN—the net amount of carbon converted to biomass. 3-PG uses a simple relationship to estimate the amount of carbon allocated below ground and a procedure based on allometric ratios—widely available for many species and situations—to determine the allocation of carbon to foliage and stems and constrain tree growth patterns. The effects of nutrition are incorporated through the carbon allocation procedure; the amount of carbon allocated below ground will increase with decreasing soil fertility. Recently acquired knowledge about the physiological factors causing decline in forest growth rates with age is used to model that decline. Changes in stem populations (self-thinning) are derived from a procedure based on the −32 power law, combined with stem growth rates. 
 
The model requires weather data as input, works on monthly time steps and has been run for periods up to 120 years, producing realistic patterns of stem growth and stem diameter increments. The time course of leaf area index is realistic for a range of soil conditions and atmospheric constraints. 3-PG can be run from remotely-sensed estimates of leaf area index coupled to weather data and basic, readily available information about soils and stand characteristics. It is being tested as a practical tool against forestry data from New South Wales, Tasmania, Victoria and New Zealand. Test results show excellent correspondence between stand growth measurements and simulated stem growth over 30 years.

title: Tree growth‐climate relationships at the northern boreal forest tree line of North America: Evaluation of potential response to increasing carbon dioxide
abstract: Tree growth at the northern limit of the range of boreal forests is primarily limited by temperature-related factors. Thus the position of this range limit, and the growth rates of trees along the northern forest border, may undergo significant change if predictions of enhanced greenhouse warming at northern latitudes are realized. In this paper we evaluate tree ring width and maximum latewood density chronologies of white spruce for three temperature-sensitive tree line sites in northern North America: in the Brooks Range, Alaska, the Franklin Mountains, Northwest Territories, and Churchill, Manitoba. The ring width data, which more strongly integrate low-frequency temperature trends than the density series, show overall enhanced growth and inferred warming during the period of anthropogenic increase in greenhouse gases. The recent growth at these sites equals or exceeds that which has occurred during earlier centuries of more clearly natural climate variability. When the ring width and density variations are estimated using temperature and precipitation data in principal components regression analysis, no substantial residual trends are detected which might require CO2 or other nutrient fertilization as an additional explanation for recent growth changes.

title: Growth enhancement due to global atmospheric change as predicted by terrestrial ecosystem models: consistent with US forest inventory data
abstract: Small reported growth enhancement factors based on analyses of forest inventory data from the eastern USA (Caspersen et al. 2000, Science, 290, 1148–1151) have been interpreted as evidence against CO2 fertilization in natural forests. We show to the contrary that growth enhancement in response to rising CO2, as found in ecosystems with experimental CO2 enrichment and implemented in terrestrial ecosystem models, is consistent with the data that have been presented within their uncertainties. Comparing forest inventory data with results of an empirical model of age-dependent biomass accumulation, we find that growth enhancement of plausible magnitude could not be detected in these data, even if it were present. Although forest regrowth due to land-use change is recognized as an important cause of carbon uptake by eastern US forests, forest inventory data do not provide a basis for eliminating environmentally induced growth enhancement as a substantial contribution to the global terrestrial carbon sink.

title: Photosynthetic responses of birch and alder saplings grown in a free air CO2 enrichment system in northern Japan
abstract: Though birch and alder are the common pioneer tree species which dominate in northeast Asia, little is known about the effects of the predicted increase in atmospheric CO2 concentrations ([CO2]) upon their photosynthesis in field conditions. To investigate this, we grew 2-year-old saplings of three Betulaceae species (Betula platyphylla var. japonica Hara, Betula maximowicziana Regel, and Alnus hirsuta Turcz) for 2 years in a free air CO2 enrichment system in northern Japan. Since the effect of high [CO2] is known to depend on soil conditions, we evaluated the responses in two soils which are widely distributed in northern Japan: infertile and immature volcanic ash (VA) soil, and fertile brown forest (BF) soil. For B. platyphylla, photosynthetic down-regulation occurred in both soils, but for B. maximowicziana, down-regulation occurred only in VA soil. The explanation is reduced nitrogen and Rubisco content in the leaf. For A. hirsuta, down-regulation occurred only in BF soil because of the accumulation of starch in foliage, which restricts CO2 diffusion inside the chloroplast. The higher photosynthetic rate of A. hirsuta in infertile VA soil could be due to the sink for photosynthates in the N2-fixing symbiont. These three species are all able to down-regulate at high [CO2]. However, it is possible that A. hirsuta would dominate in VA soil and B. maximowicziana in BF soil in the early stages of forest succession in a CO2-enhanced world.

title: Elevated CO(2) concentration affects leaf photosynthesis-nitrogen relationships in Pinus taeda over nine years in FACE.
abstract: To investigate whether long-term elevated carbon dioxide concentration () causes declines in photosynthetic enhancement and leaf nitrogen (N) owing to limited soil fertility, we measured photosynthesis, carboxylation capacity and area-based leaf nitrogen concentration (N(a)) in Pinus taeda L. growing in a long-term free-air CO(2) enrichment (FACE) facility at an N-limited site. We also determined how maximum rates of carboxylation (V(cmax)) and electron transport (J(max)) varied with N(a) under elevated [CO(2)]. In trees exposed to elevated [CO(2)] for 5 to 9 years, the slope of the relationship between leaf photosynthetic capacity (A(net-Ca)) and N(a) was significantly reduced by 37% in 1-year-old needles, whereas it was unaffected in current-year needles. The slope of the relationships of both V(cmax) and J(max) with N(a) decreased in 1-year-old needles after up to 9 years of growth in elevated [CO(2)], which was accompanied by a 15% reduction in N allocation to the carboxylating enzyme. Nitrogen fertilization (110 kg N ha(-1)) in the ninth year of exposure to elevated [CO(2)] restored the slopes of the relationships of V(cmax) and J(max) with N(a) to those of control trees (i.e., in ambient [CO(2)]). The J(max):V(cmax) ratio was unaffected by either [CO(2)] or N fertilization. Changes in the apparent allocation of N to photosynthetic components may be an important adjustment in pines exposed to elevated [CO(2)] on low-fertility sites. We conclude that fundamental relationships between photosynthesis or its component processes with N(a) may be altered in aging pine needles after more than 5 years of exposure to elevated atmospheric [CO(2)].

title: Subalpine Tree Growth, Climate, and Increasing CO_2: An Assessment of Recent Growth Trends
abstract: Five tree-ring series from foxtail pine (Pinus balfouriana), lodgepole pine (P. murrayana), and western juniper (Juniperus occidentalis) collected in the Sierra Nevada, California, were analyzed to determine if the temporal and spatial patterns of recent growth were consistent with the hypothesized CO{sub 2}-induced growth enhancement. Specifically, the author addresses the following questions: (1) can growth trends be explained solely in terms of climatic variation; (2) are recent growth trends unusual with respect to long-term growth records While the results offer no support for the hypothesized CO{sub 2} fertilization effect, they do provide insights into the response of subalpine conifers to climatic variation. Response surfaces demonstrate that precipitation during previous winter and temperature during the current summer interact in controlling growth and that the response can be nonlinear. Although maximum growth rates occur under conditions of high winter precipitation and warm summers for all three species, substantial species-to-species variation occurs in the response to these two variables.

title: Computer-Assisted Quality Control in Tree-Ring Dating and Measurement
abstract: Data quality control is especially important for tree -ring measurement series from non -arid regions. Since climatic factors are less often severely limiting to tree growth in these regions, ring series are less sensitive, and the climatic signal contained therein is weaker than in arid sites (Fritts 1976). In such areas it is very important to preserve as much as possible of the climatic information through accurate crossdating and measurement, with a minimum of "noise" introduced through errors. In determining the dating of tree -ring site collections from areas of somewhat difficult crossdating, an objective and efficient tool for locating errors or for confirming the crossdating would be very useful. Fritts describes some of the difficulties encountered in crossdating and emphasizes the importance of accurate dating for inferring climate from tree -rings, particularly since precise time definition is one of the advantageous qualities of dendrochronological data in comparison with most other paleoclimatic data. In addition, tree -ring series are occasionally found which though apparently of usable quality, fail to yield a conclusive crossdating match using the technique of skeleton plotting (Stokes and Smiley 1968), or by the widely used method of cross matching from memory. In such a series an injury or suppression of growth may prevent the dendrochronologist from following the dating inward from the date of collection. A new computer program for objectively checking tree -ring measurement series and aiding in the crossdating process is presented here.

title: Canadian historical and homogeneous temperature datasets for climate change analyses
abstract: The Canadian Historical Temperature Database (CHTD) was developed to meet the need for detailed individual station datasets and to produce an improved historical climate change database. It contains datasets of monthly mean maximum and minimum temperatures for 210 Canadian stations. Stations were selected on the basis of length of record, data completeness, and spatial distribution across the country. Records from separate stations were sometimes joined to temporally extend their series backward. Missing data gaps were estimated using highly correlated neighbour stations. Relative homogeneity was assessed using a Canadian developed technique based on regression models. Nonclimatic steps resulting from station or site relocations were identified and quantified. Data adjustments were performed for large steps (greater than 0.6°C) with or without metadata and for medium steps (0.4–0.6°C) with support. A bias in minimum temperatures was also identified and adjusted at principal stations located in eastern Canada. The bias results from a change in observing procedure in 1961 throughout the country, and it typically produces a decreasing step of about 0.6–0.8°C in the annual series in the eastern part of the country. Although also detectable in western Canada, it appears to be small there and no bias adjustments were performed in the western part of the country. Large temporal and spatial differences in data availability exist between areas north and south of 60°N latitude making national analyses possible for only the latest 50 years of data. Spatial presentation of the linear trends before and after adjustments shows overall improvement to the regional and national trends in terms of spatial consistency. The CHTD contains the best available monthly temperature data in Canada and these datasets are now available to the climate change research community.

title: Rehabilitation and analysis of Canadian daily precipitation time series
abstract: The goal of this project was to develop adjustment procedures to use daily resolution data to generate high quality time series of precipitation and to perform regional trend analyses on the resulting datasets. A total of 69 locations, most with data covering the period 1900–96 were used. Data availability in much of the Canadian Arctic was restricted to 1948–96. By using daily data, improved corrections to precipitation data, not practical with monthly data, could be implemented. For each of three rain gauge types, corrections to account for wind undercatch and evaporation were implemented. Gauge specific wetting loss corrections were applied for each rainfall event. For snowfall, ruler measurements were used throughout the time series, to minimize potential discontinuities introduced by the adoption of Nipher shielded snow gauge measurements in the mid‐1960s. Density corrections based upon coincident ruler and Nipher measurements were applied to all ruler measurements. Where necessary, records ...

title: Canadian national tree aboveground biomass equations
abstract: The estimation of aboveground biomass density (organic dry mass per unit area) is required for balancing Canadian national forest carbon budgets. Tree biomass equations are the basic tool for converting inventory plot data into biomass density. New sets of national tree biomass equations have therefore been produced from archival biomass data collected at the beginning of the 1980s through the ENergy from the FORest research program (ENFOR) of the Canadian Forest Service. Since the sampling plan was not standardized among provinces and territories, data had to be harmonized before any biomass equation could be considered at the national level. Two features characterize the new equations: estimated biomass of the compartments (foliage, branch, wood, and bark) are constrained to equal the total biomass, and dependence among error terms for the considered compartments of the same tree is taken into account in the estimates of both the model parameters and the variance prediction. The estimation method known ...

title: Natural and anthropogenic changes in atmospheric CO2 over the last 1000 years from air in Antarctic ice and firn
abstract: A record of atmospheric CO2 mixing ratios from 1006 A.D. to 1978 A.D. has been produced by analysing the air enclosed in three ice cores from Law Dome, Antarctica. The enclosed air has unparalleled age resolution and extends into recent decades, because of the high rate of snow accumulation at the ice core sites. The CO2 data overlap with the record from direct atmospheric measurements for up to 20 years. The effects of diffusion in the firn on the CO2 mixing ratio and age of the ice core air were determined by analyzing air sampled from the surface down to the bubble close-off depth. The uncertainty of the ice core CO2 mixing ratios is 1.2 ppm (1 σ). Preindustrial CO2 mixing ratios were in the range 275–284 ppm, with the lower levels during 1550–1800 A.D., probably as a result of colder global climate. Natural CO2 variations of this magnitude make it inappropriate to refer to a single preindustrial CO2 level. Major CO2 growth occurred over the industrial period except during 1935–1945 A.D. when CO2 mixing ratios stabilized or decreased slightly, probably as a result of natural variations of the carbon cycle on a decadal timescale.

title: Tests of the RCS Method for Preserving Low-Frequency Variability in Long Tree-Ring Chronologies
abstract: To preserve multicentennial length variability in annual tree -ring chronologies, the Regional Curve Standardization (RCS) method calculates anomalies from a regionally common, non -climatic age -trend function. The influence of various factors on the estimation of the regional curve (RC) and resulting RCS chronology is discussed. These factors are: the method of calculating anomalies from the age -trend function, estimation of the true pith offset, the number of series used, species composition, and site characteristics. By applying RCS to a collection of millenniumlength tree -ring data sets, the potential and limitations of the RCS method are investigated. RCS is found to be reasonably robust with respect to tested factors, suggesting the method is a suitable tool for preserving lowfrequency variance in long tree -ring chronologies.

title: Methods of Den-drochronology: Applications in the Environmental Sciences
abstract: Some Historical Background on Dendrochronology.- Primary Data.- Data Analysis.- Methods of Calibration, Verification, and Reconstruction.- Tree-Ring/Environment Interactions and Their Assessment.- Tree Rings in the Study of Future Change.

title: On the Average Value of Correlated Time Series, with Applications in Dendroclimatology and Hydrometeorology
abstract: In a number of areas of applied climatology, time series are either averaged to enhance a common underlying signal or combined to produce area averages. How well, then, does the average of a finite number (N) of time series represent the population average, and how well will a subset of series represent the N-series average? We have answered these questions by deriving formulas for 1) the correlation coefficient between the average of N time series and the average of n such series (where n is an arbitrary subset of N) and 2) the correlation between the N-series average and the population. We refer to these mean correlations as the subsample signal strength (SSS) and the expressed population signal (EPS). They may be expressed in terms of the mean inter-series correlation coefficient r as SSS ≡ (Rn,N)2 ≈ n(1 + (N − 1)r)/ N(1 + (N − 1)r), EPS ≡ RN)2 ≈ Nr/1 + (N − 1)r.Similar formulas are given relating these mean correlations to the fractional common variance which arises as a parameter in a...

title: Modeling the influence of temperature on monthly gross primary productivity of sugar maple stands.
abstract: A bottom-up and a top-down model were used to estimate the effect of temperature on monthly gross primary productivity (GPP) of sugar maple (Acer saccharum Marsh.). The bottom-up model computed canopy photosynthesis at an hourly time step from detailed physiological sub-models of leaf photosynthesis and stomatal conductance. Leaf mass per area was used as a covariable to integrate photosynthesis through the canopy. The top-down model used a radiation-use efficiency coefficient to relate canopy gross photosynthesis to absorbed photosynthetically active radiation at a monthly time step. The parameters of the top-down model were estimated from simulations with the bottom-up model. Forty single-year simulations were made using records of daily maximum and minimum temperatures from weather stations selected within the natural range of sugar maple in the province of Québec, Canada. Leaf area index was randomly varied between 4 and 10. Within a broad range of values, temperature had a minor effect on predicted monthly canopy-level GPP and its contribution to explaining the variability of GPP was low, both through its direct effect on photosynthetic processes (1.1%), and indirectly through the effect of relative humidity on stomatal conductance (4.0%). This result was unchanged when key parameters relating photosynthesis to temperature and stomatal conductance to atmospheric humidity were changed in the bottom-up model. An increase in time step from hourly to monthly resulted in a downward shift in the optimum temperature range for photosynthesis, from 30 degrees C for a leaf at saturating irradiance to 22 degrees C for the canopy at a monthly time scale.

title: Response of tree growth to a changing climate in boreal central Canada: A comparison of empirical, process-based, and hybrid modelling approaches
abstract: The impact of 2 × CO 2 driven climate change on radial growth of boreal tree species Pinus banksiana Lamb., Populus tremuloides Michx. and Picea mariana (Mill.) BSP growing in the Duck Mountain Provincial Forest of Manitoba (DMPF), Canada, is simulated using empirical and process-based model approaches. First, empirical relationships between growth and climate are developed. Stepwise multiple-regression models are conducted between tree-ring growth increments (TRGI) and monthly drought, precipitation and temperature series. Predictive skills are tested using a calibration–verification scheme. The established relationships are then transferred to climates driven by 1× and 2 × CO 2 scenarios using outputs from the Canadian second-generation coupled global climate model. Second, empirical results are contrasted with process-based projections of net primary productivity allocated to stem development (NPP s ). At the finest scale, a leaf-level model of photosynthesis is used to simulate canopy properties per species and their interaction with the variability in radiation, temperature and vapour pressure deficit. Then, a top-down plot-level model of forest productivity is used to simulate landscape-level productivity by capturing the between-stand variability in forest cover. Results show that the predicted TRGI from the empirical models account for up to 56.3% of the variance in the observed TRGI over the period 1912–1999. Under a 2 × CO 2 scenario, the predicted impact of climate change is a radial growth decline for all three species under study . However, projections obtained from the process-based model suggest that an increasing growing season length in a changing climate could counteract and potentially overwhelm the negative influence of increased drought stress. The divergence between TRGI and NPP s simulations likely resulted, among others, from assumptions about soil water holding capacity and from calibration of variables affecting gross primary productivity. An attempt was therefore made to bridge the gap between the two modelling approaches by using physiological variables as TRGI predictors. Results obtained in this manner are similar to those obtained using climate variables, and suggest that the positive effect of increasing growing season length would be counteracted by increasing summer temperatures. Notwithstanding uncertainties in these simulations (CO 2 fertilization effect, feedback from disturbance regimes, phenology of species, and uncertainties in future CO 2 emissions), a decrease in forest productivity with climate change should be considered as a plausible scenario in sustainable forest management planning of the DMPF.

title: Importance of needle age and shoot structure on canopy net photosynthesis of balsam fir (Abies balsamea): a spatially inexplicit modeling analysis.
abstract: We have developed a spatially inexplicit model of canopy photosynthesis for balsam fir (Abies balsamea (L.) Mill.) that accounts for key processes of light-shoot interaction including irradiance interception by the shoot, spatial aggregation of shoots into branches and crowns, the differential propagation of diffuse and direct light within the canopy, and an ideal representation of penumbra. Also accounted for in the model are the effects of the average radiative climate and shoot age on needle retention, light interception, and photosynthetic capacity. We used reduced versions of this model to quantify the effects of simplifying canopy representation on modeled canopy net photosynthesis. Simplifications explored were the omission of direct beam transformation into penumbral light and the use of different constant shoot properties throughout the canopy. The model was parameterized for a relatively dense balsam fir stand (leaf area index of 5.8) north of Québec City, Canada, and run using hourly meteorological data obtained at the site. The overall performance of the complete model was satisfactory, with maximum values of canopy net photosynthesis of 23 micromol (m(2) ground)(-1) s(-1) (83 mmol m(-2) h(-1)), and a near-saturation of the canopy at a photosynthetically active radiation photon flux density of about 750 micromol m(-2) s(-1) (2.7 mol m(-2) h(-1)). The omission of penumbral effects through the use of unattenuated direct (beam) radiation at all layers of the canopy, as used for broad-leaved species, reduced canopy net photosynthesis by 3.7%. Analysis of the results show that the small impact of penumbra on canopy net photosynthesis stems from the high proportion of diffuse radiation (73%) estimated from our meteorological data set; single-hour results under clear sky conditions approach theoretical bias values of about 30%. Use of mean shoot photosynthetic, light capture and light transmission properties throughout the canopy biased canopy net photosynthesis by less than 3%. However, simulations carried out based on properties of 1-year-old shoots throughout the canopy overestimated canopy net photosynthesis by 9%. Use of the shoot as our smallest functional unit was a potential source of bias because the differential absorption of direct and diffuse radiation within the shoot could not be factored into the model. Other sources of potential bias are discussed.

title: Validation of a canopy gas exchange model and derivation of a soil water modifier for transpiration for sugar maple (Acer saccharum Marsh.) using sap flow density measurements
abstract: Hourly plot-level transpiration measurements were carried out in a stand of sugar maple (Acer saccharum Marsh.) near Quebec City, Canada, during one summer using radial flow meters installed in selected trees. The measurements were used to validate transpiration estimates obtained from a multi-layer gas exchange model that included the modeling of the within-canopy radiation regime and the thermal balance of the leaves. The comparison between modeled and measured hourly transpiration showed no bias in the predictions, and an r2 value of 0.80. Because of the close coupling between transpiration and photosynthesis built in the model, these results suggest that modeled estimates of photosynthesis should also be well-related to actual rates at the stand-level. The transpiration data were also used to adjust an empirical transpiration model based on the Penman–Monteith equation in which the canopy conductance term was replaced by a function of global radiation, vapor pressure deficit and soil water depletion. This empirically-fitted model captured 85% of the variation observed in the data, including the effect of soil water depletion during a late-season drought. The soil water modifier included within this equation was compared with other soil water modifier functions obtained from the literature. The comparison highlights two difficulties in the derivation of soil water modifiers applicable outside experimental areas. The first is in the determination of rooting depth so that mass balances of soil water content can be carried out. The second is in the determination of soil physical properties so that absolute values of moisture contents can be translated into relative values of water availability.

title: Leaf nitrogen, photosynthesis, conductance and transpiration : scaling from leaves to canopies
abstract: A model is presented which solves simultaneously for leafâscale stomatal conductance, CO2 assimilation and the energy balance as a function of leaf position within canopies of wellâwatered vegetation. Fluxes and conductances were calculated separately for sunlit and shaded leaves. A linear dependence of photosynthetic capacity on leaf nitrogen content was assumed, while leaf nitrogen content and light intensity were assumed to decrease exponentially within canopies. Separate extinction coefficients were used for diffuse and direct beam radiation. An efficient Gaussian integration technique was used to compute fluxes and mean conductances for the canopy. The multilayer model synthesizes current knowledge of radiation penetration, leaf physiology and the physics of evaporation and provides insights into the response of whole canopies to multiple, interacting factors. The model was also used to explore sources of variation in the slopes of two simple parametric models (nitrogenâ and lightâuse efficiency), and to set bounds on the magnitudes of the parameters. For canopies low in total N, daily assimilation rates are â¼10% lower when leaf N is distributed uniformly than when the same total N is distributed according to the exponentially decreasing profile of absorbed radiation. However, gains are negligible for plants with high N concentrations. Canopy conductance, Gc should be calculated as Gc=AI(fslgsl+fshgsh), where I is leaf area index, fsi and fsh are the fractions of sunlit and shaded leaves at each level, and gsi and gsh are the corresponding stomatal conductances. Simple addition of conductances without this weighting causes errors in transpiration calculated using the âbigâleafâ version of the PenmanâMonteith equation. Partitioning of available energy between sensible and latent heat is very responsive to the parameter describing the sensitivity of stomata to the atmospheric humidity deficit. This parameter also affects canopy conductance, but has a relatively small impact on canopy assimilation. Simple parametric models are useful for extrapolating understanding from small to large scales, but the complexity of real ecosystems is thus subsumed in unexplained variations in parameter values. Simulations with the multilayer model show that both nitrogenâ and radiationâuse efficiencies depend on plant nutritional status and the diffuse component of incident radiation, causing a 2â to 3âfold variation in these efficiencies.

title: BOREAS in 1997: Experiment overview, scientific results, and future directions
abstract: The goal of the Boreal Ecosystem-Atmosphere Study (BOREAS) is to improve our understanding of the interactions between the boreal forest biome and the atmosphere in order to clarify their roles in global change. This overview paper describes the science background and motivations for BOREAS and the experimental design and operations of the BOREAS 1994 and BOREAS 1996 field years. The findings of the 83 papers in this journal special issue are reviewed. In section 7, important scientific results of the project to date are summarized and future research directions are identified.

title: Predicting the effects of climate change on water yield and forest production in the northeastern United States
abstract: Rapid and simultaneous changes in temperature, precipitation and the atmospheric concentration of CO2 are predicted to occur over the next century. Simple, well-validated models of ecosystem function are required to predict the effects of these changes. This paper describes an improved version of a forest carbon and water balance model (PnET-II) and the application of the model to predict stand- and regional-level effects of changes in temperature, precipitation and atmospheric CO2 concentration. PnET-II is a simple, generalized, monthly time-step model of water and carbon balances (gross and net) driven by nitrogen availability as expressed through foliar N concentration. Improvements from the original model include a complete carbon balance and improvements in the prediction of canopy phenology, as well as in the computation of canopy structure and photosynthesis. The model was parameterized and run for 4 forest/site combinations and validated against available data for water yield, gross and net carbon exchange and biomass production. The validation exercise suggests that the determination of actual water availability to stands and the occurrence or non-occurrence of soil-based water stress are critical to accurate modeling of forest net primary production (NPP) and net ecosystem production (NEP). The model was then run for the entire NewEngland/New York (USA) region using a 1 km resolution geographic information system. Predicted long-term NEP ranged from -85 to +275 g C m-2 yr-1 for the 4 forest/site combinations, and from -150 to 350 g C m-2 yr-1 for the region, with a regional average of 76 g C m-2 yr-1. A combination of increased temperature (+6*C), decreased precipitation (-15%) and increased water use efficiency (2x, due to doubling of CO2) resulted generally in increases in NPP and decreases in water yield over the region.

title: A generalized, lumped-parameter model of photosynthesis, evapotranspiration and net primary production in temperate and boreal forest ecosystems
abstract: SummaryPnET is a simple, lumped-parameter, monthlytime-step model of carbon and water balances of forests built on two principal relationships: 1) maximum photosynthetic rate is a function of foliar nitrogen concentration, and 2) stomatal conductance is a function of realized photosynthetic rate. Monthyly leaf area display and carbon and water balances are predicted by combining these with standard equations describing light attenuation in canopies and photosynthetic response to diminishing radiation intensity, along with effects of soil water stress and vapor pressure deficit (VPD). PnET has been validated against field data from 10 well-studied temperate and boreal forest ecosystems, supporting our central hypothesis that aggregation of climatic data to the monthly scale and biological data such as foliar characteristics to the ecosystem level does not cause a significant loss of information relative to long-term, mean ecosystem responses. Sensitivity analyses reveal a diversity of responses among systems to identical alterations in climatic drivers. This suggests that great care should be used in developing generalizations as to how forests will respond to a changing climate. Also critical is the degree to which the temperature responses of photosynthesis and respiration might acclimate to changes in mean temperatures at decadal time scales. An extreme climate change simulation (+3° C maximum temperature, −25% precipitation with no change in minimum temperature or radiation, direct effects of increased atmospheric CO2 ignored) suggests that major increases in water stress, and reductions in biomass production (net carbon gain) and water yield would follow such a change.

title: Population respiration a theoretical approach
abstract: It is shown that the error in an estimate of the population respiration should generally not exceed 10% if the mean body weight is used instead of the full weight distribution. A convenient mathematical formulation of the Krogh-curve is given such that the commonly used exponential temperature dependence ( Q 10 law) of the respiration rate is only a special case. A formula is derived which expresses the effect of a cyclic temperature relative to a constant temperature on the respiration of an animal, the respiration rate of which follows a Krogh-curve. This formula is shown to provide good estimates of annual population respiration both for cases where the population biomass can be considered as constant over the year and when its variation can be approximated with a sinusfunction. Our methods of estimating population respiration are shown to compare favourably with earlier shortcut methods.

title: Growth and maintenance respiration rates of aspen, black spruce and jack pine stems at northern and southern BOREAS sites.
abstract: We measured stem respiration rates during and after the 1994 growing season of three common boreal tree species at sites near the northern and southern boundaries of the closed-canopy boreal forest in central Canada. The growth respiration coefficient (r(g); carbon efflux per micro mole of carbon incorporated in structural matter) varied between 0.25 and 0.76, and was greatest for black spruce (Picea mariana (Mill.) B.S.P.), least for jack pine (Pinus banksiana Lamb.) and intermediate for trembling aspen (Populus tremuloides Michx.). There was a consistent trend for higher r(g) at northern sites than at southern sites. Maintenance respiration rates at 15 degrees C (r(m)) varied from 0.5 to 2.7 nmol C mol(-1) C(sapwood) s(-1). Values of r(m) were high at sapling-stage jack pine sites and mature black spruce sites, and low at mature trembling aspen and mature jack pine sites. We found significant relationships between annual maintenance respiration and sapwood relative growth rate and sapwood volume per unit of stem surface area that explained much of the within-stand and between-stand variability. Because of the large differences in parameter values among sites, we conclude that the use of stand-specific respiratory parameters may improve model predictions of ecosystem process models over the use of generic parameter values.

title: On the temperature dependence of soil respiration
abstract: From previously published measurements of soil respiration rate (R) and temperature (T) the goodness of fit of various R vs T relationships was evaluated. Exponential (Q 10 ) and conventional Arrhenius relationships between T and R cannot provide an unbiased estimate of respiration rate. Nor is a simple linear relationship appropriate. The relationship between R and T can, however, be accurately represented by an Arrhenius type equation where the effective activation energy for respiration varies inversely with temperature. An empirical equation is presented which yields an unbiased estimator of respiration rates over a wide range of temperatures. When combined with seasonal estimates of Gross Primary Productivity (GPP) the empirical relationship derived provides representative estimates of the seasonal cycle of net ecosystem productivity and its effects on atmospheric CO 2 (...)

title: Use of modeled photosynthesis and decomposition to describe tree growth at the northern tree line.
abstract: Growth of subarctic Scots pine (Pinus sylvestris L.) trees was investigated by a combination of process-based models and dendroecological approaches. Tree ring width indices were strongly autocorrelated and correlated with simulated photosynthetic production of the previous year and with organic matter N mineralization of the current year. An autoregressive model, with photosynthesis and N mineralization as external inputs, explained growth of the trees well. However, relationships for the period 1950-1992 differed significantly from relationships for the period 1876-1949; the slope of the regression of tree ring width index and photosynthesis was lower for the 1950-1992 period. Also, the autocorrelation structure of the data changed. First-order autocorrelation decreased and second-order autocorrelation increased from the earlier to the later period. This means that growth is becoming less sensitive to variations in photosynthetic production, whereas the relationships between growth and N mineralization are remaining fairly constant. We postulate that, although photosynthesis has increased in response to increasing CO2 concentrations, tree growth rate cannot parallel the increase in photosynthesis because potential growth rate is limited directly by temperature.

title: Carbon distribution and aboveground net primary production in aspen, jack pine, and black spruce stands in Saskatchewan and Manitoba, Canada
abstract: The objectives of this study are to (1) characterize the carbon (C) content, leaf area index, and aboveground net primary production (ANPP) for mature aspen, black spruce, and young and mature jack pine stands at the southern and northern Boreal Ecosystem-Atmosphere Study (BOREAS) areas and (2) compare net primary production and carbon allocation coefficients for the major boreal forest types of the world. Direct estimates of leaf area index, defined as one half of the total leaf surface area, range from a minimum of 1.8 for jack pine forests to a maximum of 5.6 for black spruce forests; stems comprise 5 to 15% of the total overstory plant area. In the BOREAS study, total ecosystem (vegetation plus detritus plus soil) carbon content is greatest in the black spruce forests (445,760–479,380 kg C ha−1), with 87 to 88% of the C in the soil, and is lowest in the jack pine stands (68,370–68,980 kg C ha−1) with a similar distribution of carbon in the vegetation and soil. Forest floor carbon content and mean residence time (MRT) also vary more among forest types in a study area than between study areas for a forest type; forest floor MRT range from 16 to 19 years for aspen stands to 28 to 39 years for jack pine stands. ANPP differs significantly among the mature forests at each of the BOREAS study areas, ranging from a maximum of 3490 to 3520 kg C ha−1 yr−1 for aspen stands to 1170 to 1220 kg C ha−1 yr−1 for jack pine stands. Both net primary production (NPP) and carbon allocation differ between boreal evergreen and deciduous forests in the world, suggesting global primary production models should distinguish between these two forest types. On average, 56% of NPP for boreal forests occurs as detritus and illustrates the need to better understand factors controlling aboveground and below-ground detritus production in boreal forests.

title: Ecophysiological controls on the carbon balances of three southern boreal forests
abstract: Continuous measurements of carbon exchange using the eddy covariance (EC) technique were made at three boreal forest mature sites including Southern Old Aspen (SOA), Southern Old Black Spruce (SOBS) and Southern Old Jack Pine (SOJP) in 2000. Climatic conditions were slightly warmer than normal with precipitation exceeding evapotranspiration at each site. Annual ecosystem respiration ( R) derived from daytime analyses of EC data was 1141, 815 and 52 1 g C m−2 per year and was consistently lower than nighttime EC estimates of 1193, 897 and 57 8 g C m−2 per year for SOA, SOBS and SOJP, respectively. The differences, however, were not statistically significant given the large uncertainty associated with each analytical technique. The uncertainty in annual net ecosystem productivity (NEP) was assessed by randomly simulating missing data and gap filling using simple biophysical algorithms. The uncertainty analysis supports the finding that each site was a net sink, and that differences in NEP were only significant between SOA and SOBS. The annual NEP and uncertainty for SOA, SOBS and SOJP was 122 (64–142), 35 (18–53) and 78 (61–91 ) g C m−2 per year, respectively. These relatively old growth forests represent a weak to moderate carbon sink. Despite having the shortest growing period, carbon sequestration was greatest at SOA because of its relatively large photosynthetic capacity ( Amax). At the evergreen sites, Amax was marginally larger at SOBS; however, annual carbon sequestration was smaller as a result of greater R. The evergreen sites exhibited a pronounced mid-season reduction in NEP, which was attributed to a large increase in R wh le Amax had not reached its full capacity. Non-growing season R resulted in a carbon loss of 285, 120 and 6 4 g C m−2 and accounted for 70, 80 and 46% of the summertime NEP at SOA, SOBS and SOJP, respectively. Six years of EC data at SOA indicate that carbon sequestration at boreal aspen sites may benefit from warmer climatic conditions because R is r latively conservative and photosynthesis increases in response to a longer growing period.

title: Assessing the eddy covariance technique for evaluating carbon dioxide exchange rates of ecosystems: past, present and future
abstract: The eddy covariance technique ascertains the exchange rate of CO2 across the interface between the atmosphere and a plant canopy by measuring the covariance between fluctuations in vertical wind velocity and CO2 mixing ratio. Two decades ago, the method was employed to study CO2 exchange of agricultural crops under ideal conditions during short field campaigns. During the past decade the eddy covariance method has emerged as an important tool for evaluating fluxes of carbon dioxide between terrestrial ecosystems and the atmosphere over the course of a year, and more. At present, the method is being applied in a nearly continuous mode to study carbon dioxide and water vapor exchange at over a hundred and eighty field sites, worldwide. The objective of this review is to assess the eddy covariance method as it is being applied by the global change community on increasingly longer time scales and over less than ideal surfaces. 
 
The eddy covariance method is most accurate when the atmospheric conditions (wind, temperature, humidity, CO2) are steady, the underlying vegetation is homogeneous and it is situated on flat terrain for an extended distance upwind. When the eddy covariance method is applied over natural and complex landscapes or during atmospheric conditions that vary with time, the quantification of CO2 exchange between the biosphere and atmosphere must include measurements of atmospheric storage, flux divergence and advection. 
 
Averaging CO2 flux measurements over long periods (days to year) reduces random sampling error to relatively small values. Unfortunately, data gaps are inevitable when constructing long data records. Data gaps are generally filled with values produced from statistical and empirical models to produce daily and annual sums of CO2 exchange. Filling data gaps with empirical estimates do not introduce significant bias errors because the empirical algorithms are derived from large statistical populations. On the other hand, flux measurement errors can be biased at night when winds are light and intermittent. Nighttime bias errors tend to produce an underestimate in the measurement of ecosystem respiration. 
 
Despite the sources of errors associated with long-term eddy flux measurements, many investigators are producing defensible estimates of annual carbon exchange. When measurements come from nearly ideal sites the error bound on the net annual exchange of CO2 is less than ±50 g C m−2 yr−1. Additional confidence in long-term measurements is growing because investigators are producing values of net ecosystem productivity that are converging with independent values produced by measuring changes in biomass and soil carbon, as long as the biomass inventory studies are conducted over multiple years.

title: FLUXNET: A New Tool to Study the Temporal and Spatial Variability of Ecosystem-Scale Carbon Dioxide, Water Vapor, and Energy Flux Densities
abstract: FLUXNET is a global network of micrometeorological flux measurement sites that measure the exchanges of carbon dioxide, water vapor, and energy between the biosphere and atmosphere. At present over 140 sites are operating on a long-term and continuous basis. Vegetation under study includes temperate conifer and broadleaved (deciduous and evergreen) forests, tropical and boreal forests, crops, grasslands, chaparral, wetlands, and tundra. Sites exist on five continents and their latitudinal distribution ranges from 70°N to 30°S. FLUXNET has several primary functions. First, it provides infrastructure for compiling, archiving, and distributing carbon, water, and energy flux measurement, and meteorological, plant, and soil data to the science community. (Data and site information are available online at the FLUXNET Web site, http://www-eosdis.ornl.gov/FLUXNET/.) Second, the project supports calibration and flux intercomparison activities. This activity ensures that data from the regional networks are intercomparable. And third, FLUXNET supports the synthesis, discussion, and communication of ideas and data by supporting project scientists, workshops, and visiting scientists. The overarching goal is to provide information for validating computations of net primary productivity, evaporation, and energy absorption that are being generated by sensors mounted on the NASA Terra satellite. Data being compiled by FLUXNET are being used to quantify and compare magnitudes and dynamics of annual ecosystem carbon and water balances, to quantify the response of stand-scale carbon dioxide and water vapor flux densities to controlling biotic and abiotic factors, and to validate a hierarchy of soil–plant–atmosphere trace gas exchange models. Findings so far include 1) net CO 2 exchange of temperate broadleaved forests increases by about 5.7 g C m −2 day −1 for each additional day that the growing season is extended; 2) the sensitivity of net ecosystem CO 2 exchange to sunlight doubles if the sky is cloudy rather than clear; 3) the spectrum of CO 2 flux density exhibits peaks at timescales of days, weeks, and years, and a spectral gap exists at the month timescale; 4) the optimal temperature of net CO 2 exchange varies with mean summer temperature; and 5) stand age affects carbon dioxide and water vapor flux densities.

title: Analysis of historic jack pine budworm outbreaks in the Prairie provinces of Canada
abstract: The annual Forest Insect and Disease Survey reports of the Canadian Forestry Service were used to develop a jack pine budworm (Choristoneurapinus Freeman) defoliation severity index for a 50-year s...

title: Effects of seasonal and interannual climate variability on net ecosystem productivity of boreal deciduous and conifer forests
abstract: The response of net ecosystem productivity (NEP) and evaporation in a boreal aspen (Populus tremuloides Michx.) forest and a black spruce (Picea mariana (Mill.) BSP) forest in Canada was compared u...

title: Changes in net ecosystem productivity of boreal black spruce stands in response to changes in temperature at diurnal and seasonal time scales.
abstract: Net ecosystem productivity (NEP) of boreal coniferous forests is believed to rise with climate warming, thereby offsetting some of the rise in atmospheric CO(2) concentration (C(a)) by which warming is caused. However, the response of conifer NEP to warming may vary seasonally, with rises in spring and declines in summer. To gain more insight into this response, we compared changes in CO(2) exchange measured by eddy covariance and simulated by the ecosystem process model ecosys under rising mean annual air temperatures (T(a)) during 2004-2006 at black spruce stands in Saskatchewan, Manitoba and Quebec. Hourly net CO(2) uptake was found to rise with warming at T(a) < 15 degrees C and to decline with warming at T(a) > 20 degrees C. As mean annual T(a) rose from 2004 to 2006, increases in net CO(2) uptake with warming at lower T(a) were greater than declines with warming at higher T(a) so that annual gross primary productivity and hence NEP increased. Increases in net CO(2) uptake measured at lower T(a) were explained in the model by earlier recovery of photosynthetic capacity in spring, and by increases in carboxylation activity, using parameters for the Arrhenius temperature functions of key carboxylation processes derived from independent experiments. Declines in net CO(2) uptake measured at higher T(a) were explained in the model by sharp declines in mid-afternoon canopy stomatal conductance (g(c)) under higher vapor pressure deficits (D). These declines were modeled from a hydraulic constraint to water uptake imposed by low axial conductivity of conifer roots and boles that forced declines in canopy water potential (psi(c)), and hence in g(c) under higher D when equilibrating water uptake with transpiration. In a model sensitivity study, the contrasting responses of net CO(2) uptake to specified rises in T(a) caused annual NEP of black spruce in the model to rise with increases in T(a) of up to 6 degrees C, but to decline with further increases at mid-continental sites with lower precipitation. However, these contrasting responses to warming also indicate that rises in NEP with climate warming would depend on the seasonality (spring versus summer) as well as the magnitude of rises in T(a).

title: A long‐term record of carbon exchange in a boreal black spruce forest: means, responses to interannual variability, and decadal trends
abstract: We present a decadal (1994-2004) record of carbon dioxide flux in a 160-year-old black spruce forest/veneer bog complex in central Manitoba, Canada. The ecosystem shifted from a source (+41 g Cm -2, 1995) to a sink (-21 g C m -2, 2004) of CO 2 over the decade, with an average net carbon balance near zero. Annual mean temperatures increased 1-2° during the period, consistent with the decadal trend across the North American boreal biome. We found that ecosystem carbon exchange responded strongly to air temperature, moisture status, potential evapotranspiration, and summertime solar radiation. The seasonal cycle of ecosystem respiration significantly lagged that of photosynthesis, limited by the rate of soil thaw and the slow drainage of the soil column. Factors acting over long time scales, especially water table depth, strongly influenced the carbon budget on annual time scales. Net uptake was enhanced and respiration inhibited by multiple years of rainfall in excess of evaporative demand. Contrary to expectations, we observed no correlation between longer growing seasons and net uptake, possibly because of offsetting increases in ecosystem respiration. The results indicate that the interactions between soil thaw and water table depth provide critical controls on carbon exchange in boreal forests underlain by peat, on seasonal to decadal time scales, and these factors must be simulated in terrestrial biosphere models to predict response of these regions to future climate.

title: Variations in northern vegetation activity inferred from satellite data of vegetation index during 1981 to 1999
abstract: The northern high latitudes have warmed by about 0.8°C since the early 1970s, but not all areas have warmed uniformly [Hansen et al., 1999]. There is warming in most of Eurasia, but the warming rate in the United States is smaller than in most of the world, and a slight cooling is observed in the eastern United States over the past 50 years. These changes beg the question, can we detect the biotic response to temperature changes? Here we present results from analyses of a recently developed satellite-sensed normalized difference vegetation index (NDVI) data set for the period July 1981 to December 1999: (1) About 61% of the total vegetated area between 40°N and 70°N in Eurasia shows a persistent increase in growing season NDVI over a broad contiguous swath of land from central Europe through Siberia to the Aldan plateau, where almost 58% (7.3×106 km2) is forests and woodlands; North America, in comparison, shows a fragmented pattern of change in smaller areas notable only in the forests of the southeast and grasslands of the upper Midwest, (2) A larger increase in growing season NDVI magnitude (12% versus 8%) and a longer active growing season (18 versus 12 days) brought about by an early spring and delayed autumn are observed in Eurasia relative to North America, (3) NDVI decreases are observed in parts of Alaska, boreal Canada, and northeastern Asia, possibly due to temperature-induced drought as these regions experienced pronounced warming without a concurrent increase in rainfall [Barber et al., 2000]. We argue that these changes in NDVI reflect changes in biological activity. Statistical analyses indicate that there is a statistically meaningful relation between changes in NDVI and land surface temperature for vegetated areas between 40°N and 70°N. That is, the temporal changes and continental differences in NDVI are consistent with ground-based measurements of temperature, an important determinant of biological activity. Together, these results suggest a photosynthetically vigorous Eurasia relative to North America during the past 2 decades, possibly driven by temperature and precipitation patterns. Our results are in broad agreement with a recent comparative analysis of 1980s and 1990s boreal and temperate forest inventory data [United Nations, 2000].

title: Increased activity of northern vegetation inferred from atmospheric CO2 measurements
abstract: THROUGHOUT the Northern Hemisphere the concentration of atmospheric carbon dioxide rises in winter and declines in summer, mainly in response to the seasonal growth in land vegetation1–4. In the far north the amplitude of the seasonal cycle, peak to trough, is between 15 and 20 parts per million by volume5. The annual amplitude diminishes southwards to about 3 p.p.m. near the Equator, owing to the diminishing seasonally of plant activity towards the tropics. In spite of atmospheric mixing processes, enough spatial variability is retained in the seasonal cycle of CO2 to reveal considerable regional detail in seasonal plant activity6. Here we report that the annual amplitude of the seasonal CO2 cycle has increased by 20%, as measured in Hawaii, and by 40% in the Arctic, since the early 1960s. These increases are accompanied by phase advances of about 7 days during the declining phase of the cycle, suggesting a lengthening of the growing season. In addition, the annual amplitudes show maxima which appear to reflect a sensitivity to global warming episodes that peaked in 1981 and 1990. We propose that the amplitude increases reflect increasing assimilation of CO2 by land plants in response to climate changes accompanying recent rapid increases in temperature.

title: Height growth response of tree line black spruce to recent climate warming across the forest‐tundra of eastern Canada
abstract: The northward expansion of the boreal forest vegetation zone is generally predicted under a warmer doubled CO 2 , but the delay associated with vegetation development processes often has been overlooked. In the subarctic forest-tundra of northern Quebec, reforestation of tundra uplands appears currently limited by the poor reproductive capacity of shrubby black spruce ( Picea mariana ), and the development of erect stems through accelerated height growth should be the first registered response to 20th century climate warming. The subarctic forest-tundra is characterized by small- and large-scale heterogeneity in topography, vegetation structure and climate. This spatial heterogeneity, added to the complexity of tree growth‐climate relationships, can cause various growth responses of subarctic tree line black spruce to 20th century climate change. 2 Twenty spruce populations at subarctic tree lines and seven isolated clones at the species limit were sampled along a > 300-km latitudinal transect from the southern forest-tundra to the shrub tundra. Height growth patterns of black spruce at tree line and above tree line were examined (i) over their life span, using dendrochronological dating of stem cross-sections, and (ii) for the recent decades, using leader shoot elongation measurements. Indexed elongation chronologies were compared with regional climate data. 3 Height growth of tree line trees generally decreased with increasing latitude. However, tree line trees in the northern forest-tundra have experienced an acceleration of height growth since the 1970s, with their growth comparable to that of trees in the southern forest-tundra. Height growth response of spruce trees appeared increasingly delayed from the northern forest-tundra to the species limit. Above the subarctic tree line, windexposed conditions obscured the decrease in height growth with latitude observed for tree line trees. 4 Leader shoot elongation of spruce trees established on tundra hilltops appeared more controlled by summer heat sums than those at tree line all over the forest-tundra, except at the arctic tree line. Winter precipitation also was linked to leader shoot elongation in some forest-tundra sites. The increasing snow cover associated with recent warming appeared to have reduced the shoot elongation of spruce at forest margins showing the steepest slopes, hence subjected to snow overloading. 5 In the northern forest-tundra sites, the recent increase in height growth and positive trend in leader shoot elongation, consistent with a 1990s’ increase in heat sums, point to the development of spruce krummholz into erect growth forms. In the southern foresttundra, reforestation of tundra hilltops and northward expansion of the boreal forest predicted under doubled CO 2 conditions could be delayed, as suggested by suppressed height growth of spruce above tree line.

title: Trends in recent temperature and radial tree growth spanning 2000 years across northwest Eurasia
abstract: This paper describes variability in trends of annual tree growth at several locations in the high latitudes of Eurasia, providing a wide regional comparison over a 2000-year period. The study focuses on the nature of local and widespread tree-growth responses to recent warming seen in instrumental observations, available in northern regions for periods ranging from decades to a century. Instrumental temperature data demonstrate differences in seasonal scale of Eurasian warming and the complexity and spatial diversity of tree-growing-season trends in recent decades. A set of long tree-ring chronologies provides empirical evidence of association between inter-annual tree growth and local, primarily summer, temperature variability at each location. These data show no evidence of a recent breakdown in this association as has been found at other high-latitude Northern Hemisphere locations. Using Kendall's concordance, we quantify the time-dependent relationship between growth trends of the long chronologies as a group. This provides strong evidence that the extent of recent widespread warming across northwest Eurasia, with respect to 100- to 200-year trends, is unprecedented in the last 2000 years. An equivalent analysis of simulated temperatures using the HadCM3 model fails to show a similar increase in concordance expected as a consequence of anthropogenic forcing.

title: Summer Moisture and Wildfire Risks across Canada
abstract: The Fire Weather Index System has been in use across Canada for the past 30 years in the daily operations of fire management agencies. As part of this system, the Drought Code (DC) was developed to act as a daily index of water stored in the soil. A major obstacle to the completion of climate risk analyses on the DC is that lengthy series of daily temperature and precipitation are not available for large portions of the circumboreal forest. Here the authors present a methodological modification to the daily DC to allow its approximation using monthly data. This new Monthly Drought Code (MDC) still retains its ability to capture moisture trends in deep organic layers. On the basis of high-resolution temperature and precipitation data, an analysis of summer moisture availability across Canada over 1901–2002 is presented. The driest periods on record were from the 1920s to the early 1960s, with the driest years being 1955, 1958, and 1961. The wettest period was from the mid-1960s to the 1980s. For t...

title: On the Cause of the 1930s Dust Bowl
abstract: During the 1930s, the United States experienced one of the most devastating droughts of the past century. The drought affected almost two-thirds of the country and parts of Mexico and Canada and was infamous for the numerous dust storms that occurred in the southern Great Plains. In this study, we present model results that indicate that the drought was caused by anomalous tropical sea surface temperatures during that decade and that interactions between the atmosphere and the land surface increased its severity. We also contrast the 1930s drought with other North American droughts of the 20th century.

title: Botany: Constraints to growth of boreal forests
abstract: Understanding how the growth of trees at high latitudes in boreal forest is controlled is important for projections of global carbon sequestration and timber production in relation to climate change. Is stem growth of boreal forest trees constrained by the length of the growing season when stem cambial cells divide, or by the length of the period when resources can be captured? In both cases, the timing of the thaw in the spring is critical: neither cambial cell division nor uptake of nutrients and carbon dioxide can occur while the soil is frozen. Here we argue, on the basis of long-term observations made in northern Saskatchewan and Sweden, that the time between the spring thaw and the autumn freeze determines the amount of annual tree growth, mainly through temperature effects on carbon-dioxide uptake in spring and on nutrient availability and uptake during summer, rather than on cambial cell division.

title: Nitrogen limitation constrains sustainability of ecosystem response to CO2
abstract: Enhanced plant biomass accumulation in response to elevated atmospheric CO2 concentration could dampen the future rate of increase in CO2 levels and associated climate warming. However, it is unknown whether CO2-induced stimulation of plant growth and biomass accumulation will be sustained or whether limited nitrogen (N) availability constrains greater plant growth in a CO2-enriched world. Here we show, after a six-year field study of perennial grassland species grown under ambient and elevated levels of CO2 and N, that low availability of N progressively suppresses the positive response of plant biomass to elevated CO2. Initially, the stimulation of total plant biomass by elevated CO2 was no greater at enriched than at ambient N supply. After four to six years, however, elevated CO2 stimulated plant biomass much less under ambient than enriched N supply. This response was consistent with the temporally divergent effects of elevated CO2 on soil and plant N dynamics at differing levels of N supply. Our results indicate that variability in availability of soil N and deposition of atmospheric N are both likely to influence the response of plant biomass accumulation to elevated atmospheric CO2. Given that limitations to productivity resulting from the insufficient availability of N are widespread in both unmanaged and managed vegetation, soil N supply is probably an important constraint on global terrestrial responses to elevated CO2.

title: Nitrogen availability in soil and forest floor of contrasting types of boreal mixedwood forests
abstract: Boreal mixedwood forests with varying proportions of coniferous and deciduous species are found throughout the North American continent. Maintenance of a deciduous component within boreal forests i...

title: The Contrasting Effects of Aspen and Jack Pine on Soil Nutritional Properties Depend on Parent Material
abstract: A bstractThe influence of forest stand composition on soil was investigated by comparing the forest floor (FH) and upper mineral soil (0–20 cm) nutritional properties of jack pine and aspen stands on two soil types of contrasting fertility, a coarse-textured and a fine-textured deposit, in a replicated design. The studied tree species are pioneers that are found after major disturbances in the southern boreal forest of western Quebec and that differ in their nutrient requirements but not in their growth rate. Soil organic matter as well as total and available N, P, K, Ca, Mg contents were determined and the relationships with nutrient accumulation in tree biomass were studied. On both soil types a greater total and available nutrient accumulation in the forest floor layer was observed in aspen than in jack pine whereas such differences between stand types could not be detected in the mineral soil. Differences in FH nutrient content between stand types were larger on coarse deposits than on fine-textured soils. These results support the hypothesis that tree species with greater nutrient requirements cause an enrichment of the surface soil at least in the short term. The modulation of tree species effect by soil type was contrary to the pattern observed in other studies since a greater expression of this effect was observed on poorer soils. Differences in soil nutrient content were related to levels of organic matter accumulation.

title: On the long‐term context for late twentieth century warming
abstract: Previous tree-ring–based Northern Hemisphere temperature reconstructions portray a varying amplitude range between the ‘‘Medieval Warm Period’’ (MWP), ‘‘Little Ice Age’’ (LIA) and present. We describe a new reconstruction, developed using largely different methodologies and additional new data compared to previous efforts. Unlike earlier studies, we quantify differences between more traditional (STD) and Regional Curve Standardization (RCS) methodologies, concluding that RCS is superior for retention of low-frequency trends. Continental North American versus Eurasian RCS series developed prior to merging to the hemispheric scale cohere surprisingly well, suggesting common forcing, although there are notable deviations (e.g., fifteenth to sixteenth century). Results indicate clear MWP (warm), LIA (cool), and recent (warm) episodes. Direct interpretation of the RCS reconstruction suggests that MWP temperatures were nearly 0.7C cooler than in the late twentieth century, with an amplitude difference of 1.14C from the coldest (1600–1609) to warmest (1937–1946) decades. However, we advise caution with this analysis. Although we conclude, as found elsewhere, that recent warming has been substantial relative to natural fluctuations of the past millennium, we also note that owing to the spatially heterogeneous nature of the MWP, and its different timing within different regions, present palaeoclimatic methodologies will likely ‘‘flatten out’’ estimates for this period relative to twentieth century warming, which expresses a more homogenous global ‘‘fingerprint.’’ Therefore we stress that presently available paleoclimatic reconstructions are inadequate for making specific inferences, at hemispheric scales, about MWP warmth relative to the present anthropogenic period and that such comparisons can only still be made at the local/regional scale.

title: Synoptic-Scale Atmospheric Circulation and Boreal Canada Summer Drought Variability of the Past Three Centuries
abstract: Five independent multicentury reconstructions of the July Canadian Drought Code and one reconstruction of the mean July–August temperature were developed using a network of 120 well-replicated tree-ring chronologies covering the area of the eastern Boreal Plains to the eastern Boreal Shield of Canada. The reconstructions were performed using 54 time-varying reconstruction submodels that explained up to 50% of the regional drought variance during the period of 1919–84. Spatial correlation fields on the six reconstructions revealed that the meridional component of the climate system from central to eastern Canada increased since the mid–nineteenth century. The most obvious change was observed in the decadal scale of variability. Using 500-hPa geopotential height and wind composites, this zonal to meridional transition was interpreted as a response to an amplification of long waves flowing over the eastern North Pacific into boreal Canada, from approximately 1851 to 1940. Composites with NOAA Extend...

title: The tree-ring record of drought on the Canadian Prairies
abstract: Ring-width data from 138 sites in the Canadian Prairie Provinces and adjacent regions are used to estimate summer drought severity during the past several hundred years. The network was divided into five regional groups based on geography, tree species, and length of record: the eastern Rockies, northern Saskatchewan, central Manitoba, southern Manitoba, and northwestern Ontario. Regional tree-ring records are primarily related to summer moisture and drought conditions, and are less responsive to droughts caused by deficits in winter precipitation. These summer-sensitive data are not linearly related to major modes of climate variability, including ENSO and the Pacific decadal oscillation (PDO), which primarily affect the climate of western Canada during winter. Extended drought records inferred from tree rings indicate that drought on the Canadian Prairies has exhibited considerable spatial heterogeneity over the last several centuries. For northern Saskatchewan and northwestern Ontario, intervals of persistently low tree growth during the twentieth century were just as long as or longer than low-growth intervals in the eighteenth or nineteenth centuries. Longer records from southern Alberta suggest that the most intense dry spell in that area during the last 500 yr occurred during the 1720s. At the eastern side of the prairies, the longest dry event is centered around 1700 and may coincide with low lake stands in Manitoba, Minnesota, and North Dakota. Although the Canadian Prairies were dry at times during the 1500s, there is no regional analog to the sixteenth-century ‘‘megadroughts’’ that affected much of the western United States and northern Mexico.