Frank Berninger

The human footprint in the carbon cycle of temperate and boreal forests

Temperate and boreal forests in the Northern Hemisphere cover an area of about 2 × 107 square kilometres and act as a substantial carbon sink (0.6–0.7 petagrams of carbon per year). Although forest expansion following agricultural abandonment is certainly responsible for an important fraction of this carbon sink activity, the additional effects on the carbon balance of established forests of increased atmospheric carbon dioxide, increasing temperatures, changes in management practices and nitrogen deposition are difficult to disentangle, despite an extensive network of measurement stations. The relevance of this measurement effort has also been questioned, because spot measurements fail to take into account the role of disturbances, either natural (fire, pests, windstorms) or anthropogenic (forest harvesting). Here we show that the temporal dynamics following stand-replacing disturbances do indeed account for a very large fraction of the overall variability in forest carbon sequestration. After the confounding effects of disturbance have been factored out, however, forest net carbon sequestration is found to be overwhelmingly driven by nitrogen deposition, largely the result of anthropogenic activities. The effect is always positive over the range of nitrogen deposition covered by currently available data sets, casting doubts on the risk of widespread ecosystem nitrogen saturation under natural conditions. The results demonstrate that mankind is ultimately controlling the carbon balance of temperate and boreal forests, either directly (through forest management) or indirectly (through nitrogen deposition).

Response of Forest Trees to Increased Atmospheric CO 2

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.

Drought responses of Eucalyptus microtheca provenances depend on seasonality of rainfall in their place of origin

We exposed seedlings of 12 Eucalyptus microtheca F. Muell. provenances to well-watered and water-stressed growing conditions in a greenhouse experiment and investigated the effects of drought on various plant properties in the provenances. We found significant variation in total biomass, height, root mass/foliage area ratio,foliage area/stem cross sectional area ratio, specific leaf area (SLA), water-use efficiency (WUE) and carbon isotope composition (d 13 C) among the provenances. The observed inter-provenance variation was more pronounced in the water-stressed treatment than in the well-watered one. Drought increased root mass/foliage area ratio, foliage area/stem cross sectional area ratio, WUE, d 13 C and decreased total biomass, height, transpiration and SLA. We also analysed relationships between plant properties and climate of native habitats of the provenances and found that most properties were strongly correlated with mean driest quarter rainfall. The correlation was positive for total biomass, height, transpiration and SLA and negative for root mass/foliage area ratio, foliage area/stem cross sectional area ratio, WUE and d 13 C. Finally, we evaluated the intra-specific variation in foliage area/stem cross sectional area ratio in the context of tree hydraulic architecture: provenances from dry areas and trees grown under drought stress had more foliage per stem area ratio. However, their transpiration and the length of their hydraulic pathway were smaller and therefore the root to leaf water potential gradient might be smaller in these trees.

Effect of thinning on surface fluxes in a boreal forest

[1] Thinning is a routine forest management operation that changes tree spacing, number, and size distribution and affects the material flows between vegetation and the atmosphere. Here, using direct micrometeorological ecosystem-scale measurements, we show that in a boreal pine forest, thinning decreases the deposition velocities of fine particles as expected but does not reduce the carbon sink, water vapor flux, or ozone deposition. The thinning decreased the all-sided leaf area index from 8 to 6, and we suggest that the redistribution of sources and sinks within the ecosystem compensated for this reduction in foliage area. In the case of water vapor and O 3 , changes in light penetration and among-tree competition seem to increase individual transpiration rates and lead to larger stomatal apertures, thus enhancing also O 3 deposition. In the case of CO 2 , increased ground vegetation assimilation and decreased autotrophic respiration seem to cancel out opposite changes in canopy assimilation and heterotrophic respiration. Current soil-vegetation-atmosphere transfer models should be able to reproduce these observations.

Biotic, Abiotic, and Management Controls on the Net Ecosystem CO2 Exchange of European Mountain Grassland Ecosystems

The net ecosystem carbon dioxide (CO2) exchange (NEE) of nine European mountain grassland ecosystems was measured during 2002–2004 using the eddy covariance method. Overall, the availability of photosynthetically active radiation (PPFD) was the single most important abiotic influence factor for NEE. Its role changed markedly during the course of the season, PPFD being a better predictor for NEE during periods favorable for CO2 uptake, which was spring and autumn for the sites characterized by summer droughts (southern sites) and (peak) summer for the Alpine and northern study sites. This general pattern was interrupted by grassland management practices, that is, mowing and grazing, when the variability in NEE explained by PPFD decreased in concert with the amount of aboveground biomass (BMag). Temperature was the abiotic influence factor that explained most of the variability in ecosystem respiration at the Alpine and northern study sites, but not at the southern sites characterized by a pronounced summer drought, where soil water availability and the amount of aboveground biomass were more or equally important. The amount of assimilating plant area was the single most important biotic variable determining the maximum ecosystem carbon uptake potential, that is, the NEE at saturating PPFD. Good correspondence, in terms of the magnitude of NEE, was observed with many (semi-) natural grasslands around the world, but not with grasslands sown on fertile soils in lowland locations, which exhibited higher maximum carbon gains at lower respiratory costs. It is concluded that, through triggering rapid changes in the amount and area of the aboveground plant matter, the timing and frequency of land management practices is crucial for the short-term sensitivity of the NEE of the investigated mountain grassland ecosystems to climatic drivers.

Photosynthetic responses of Populus przewalski subjected to drought stress

Cuttings of P. przewalski were exposed to two different watering regimes which were watered to 100 and 25 % of field capacity (WW and WS, respectively). Drought stress not only significantly decreased net photosynthetic rate (PN), transpiration rate (E), stomatal conductance (gs), efficiency of photosystem 2 (PS2) (Fv/Fm and yield), and increased intrinsic water use efficiency (WUEi) under controlled optimal conditions, but also altered the diurnal changes of gas exchange, chlorophyll fluorescence, and WUEi. On the other hand, WS also affected the PN-photosynthetically active radiation (PAR) response curve. Under drought stress, PN peak appeared earlier (at about 10:30 of local time) than under WW condition (at about 12:30). At midday, there was a depression in PN for WS plants, but not for WW plants, and it could be caused by the whole microclimate, especially high temperature, low relative humidity, and high PAR. There were stomatal and non-stomatal limitations to photosynthesis. Stomatal limitation dominated in the morning, and low PN at midday was caused by both stomatal and non-stomatal limitations, whereas non-stomatal limitation dominated in the afternoon. In addition, drought stress also increased compensation irradiance and dark respiration rate, and decreased saturation irradiance and maximum net photosynthetic rate. Thus drought stress decreased plant assimilation and increased dissimilation through affected gas exchange, the diurnal pattern of gas exchange, and photosynthesis-PAR response curve, thereby reducing plant growth and productivity.

Structural adaptation rather than water conservation was observed in Scots pine over a range of wet to dry climates

Abstract We investigated the functional and structural responses of Scots pine to climate and estimated the importance of the genotype on the traits studied. We analysed 13C isotope discrimination (Δ13C) of various provenances in a common garden experiment and gas exchange characteristics for provenances growing in their natural environment. No clear climatic trend was found in the foliar Δ13C values of common garden trees. Similar results were obtained from estimation of λ (a largely VPD, temperature and light independent measure of intrinsic water use efficiency) from the gas exchange data. The ratio of needle mass to unit stem area and branch area to stem area increased towards south in both experiments and hence, seemed to be genetic. Trees from drier and warmer conditions seemed not to have either lower needle mass or higher intrinsic water use efficiency compared to northern latitudes.

Fluxes of carbon dioxide and water vapour over Scots pine forest and clearing

The carbon dioxide and water vapour fluxes were measured by the eddy covariance (EC) technique from July to September 2000 at two closely located sites in southern Finland: over a 38-year-old pine forest and over a 5-year-old forest clearing. The night-time respiration was of the same magnitude at both sites. At day-time the pine forest was a strong sink but the clearing close to CO2 balance, indicating that CO2 uptake of ground vegetation over the clearing balanced the release from the soil. The shoot scale gas exchange measurements in combination with process-based modelling were used to evaluate the measured CO2 exchange of the forest ecosystem. The forest CO2 exchange was explained by soil respiration and photosynthesis of forest canopy, while the contribution of understory and ground vegetation CO2 exchange could be neglected. During the study period the forest was a net sink of CO2 and the clearing a source. The daily average uptake of CO2 by the forest was −2.4 and − 1.7 gm −2 per day in July–August and September periods, respectively; and average release by the clearing 4.0 and 2.5 g m −2 per day during the same periods. This shows that carbon losses 5 years after clear-cutting are substantial. The evapotranspiration (ET) was higher over the forest compared to clearing as a result of transpiration from the forest canopy. The difference in ET was small during the July–August period when precipitation frequently occurred.

Growth, allocation and leaf gas exchanges of hybrid poplar plants in their establishment phase on previously forested sites: effect of different vegetation management techniques

The effect of different vegetation control methods (mowing and cultivation between plantation rows, herbicide application and cover plant sowing) on hybrid poplar (P. maximowiczii × balsamifera) growth, biomass allocation and leaf carbon assimilation was investigated in two plantations (1- and 2-year-old) established in previously forested sites of south-eastern Québec. Any vegetation control treatment applied the same year in which the plantation was established did not have an effect on hybrid poplar aboveground growth. However significant differences among treatments were observed belowground, where the removal of the competing vegetation at the tree base increased the fine root:leaf biomass ratio of plants, thus probably facilitating their establishment. In contrast, 2-year-old plants grew better when treated with herbicides, but no positive effect of the mechanical treatments was detected. In both sites, trees growing on herbicide-treated plots showed considerably higher leaf carbon assimilation and leaf N concentration which were both strongly correlated. We conclude that a strong vegetation competition for nutrients takes effect on hybrid poplar plantations on previously forested sites since there was no water shortage for any treatment during the study period.RésuméL’effet de différentes méthodes de contrôle de la végétation compétitrice (fauchage et hersage mécanisé, herbicide, semence de plantes de couverture) a été étudié sur la croissance, l’allocation de biomasse et l’assimilation de carbone du peuplier hybride (P. maximowiczii × balsamifera). Les mesures ont été effectuées sur des individus provenant de deux plantations localisées sur des anciennes terres boisés dans le sud-est du Québec et établies la même année ou l’année précédant l’étude. Lorsqu’elles sont appliquées la même année que l’installation de la plantation, aucune des méthodes de contrôle de la végétation n’a eu d’effets sur la croissance aérienne des individus. Cependant, l’élimination de la végétation autour de la base des peupliers a affecté la partie racinaire des individus en augmentant le ratio de biomasse des racines fines:biomasse foliaire ce qui a probablement favorisé leur installation. Par contre, dans la plantation établie l’année antérieure, une plus forte croissance en diamètre et en hauteur a été observée chez les peupliers traités avec des herbicides alors que les traitements mécaniques n’ont encore une fois pas eu d’effet sur les individus. Dans les deux plantations, les arbres traités avec des herbicides présentaient une meilleure capacité d’assimilation de carbone et une meilleure teneure en N foliaire. Étant donné qu’aucune limitation hydrique n’a été mise en évidence pendant la période de l’étude, nous concluons qu’une forte compétition pour les éléments nutritifs existe dans les jeunes plantations de peupliers hybrides établies sur des anciennes terres boisées.

Modeling 13C discrimination in tree rings

Annual variations from 1877 to 1995 in tree-ring α-cellulose 13C/12C isotopic ratios for four subarctic Pinus sylvestris trees were determined, and, in conjunction with a recent record of atmospheric 13CO2/12CO2 ratios, the historical pattern of photosynthetic isotope discrimination, Δ13C, was evaluated. Year-to-year variability in Δ13C has been as much as 1.5‰ with the period 1900–1920 showing an extended period of unusually high photosynthetic discriminations. The summers during these years were, on average, unusually cold. Since 1920 a long term trend of increasing Δ13C of ∼0.016‰yr−1 is inferred. We compared measured Δ13C with those predicted on the basis of the theoretical relationship between Δ13C and the ratio of substomatal to ambient CO2 concentration, Ci/Ca using mechanistic equations for chloroplast biochemistry coupled with a stomatal conductance model. Two variations of a nonlinear optimal-regulation stomatal conductance model were compared. Although both models were based on the assumption that stomata serve to minimize the average transpiration rate for a given average rate of CO2 assimilation, one version of the model incorporated reductions in stomatal conductance in response to recent increases in atmospheric CO2 concentrations and the other did not. The CO2 sensitive stomatal model failed to describe the long-term increase in 13C discrimination, especially after 1950. The insensitive model gave good agreement, suggesting that an observed increase in subarctic Pinus sylvestris Δ13C since 1920 is attributable to recent increases in atmospheric CO2 concentrations with subsequent increases in the ratio of substomatal to ambient CO2 concentrations. The model was also capable of accounting for high frequency (year-to-year) variations in Δ13C, these differences being attributable to year-to-year fluctuations in the average leaf-to-air vapor pressure difference affecting stomatal conductance and hence Ci/Ca.