Temperate and boreal forests in the Northern Hemisphere cover an area of about 2 x 10(7) 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).

Summary• Intra-annual radial growth rates and durations in trees are reported to differ greatly in relation to species, site and environmental conditions. However, very similar dynamics of cambial activity and wood formation are observed in temperate and boreal zones.• Here, we compared weekly xylem cell production and variation in stem circumference in the main northern hemisphere conifer species (genera Picea , Pinus , Abies and Larix ) from 1996 to 2003. Dynamics of radial growth were modeled with a Gompertz function, defining the upper asymptote ( A ), x -axis placement ( β ) and rate of change ( κ ).• A strong linear relationship was found between the constants β and κ for both types of analysis. The slope of the linear regression, which corresponds to the time at which maximum growth rate occurred, appeared to converge towards the summer solstice.• The maximum growth rate occurred around the time of maximum day length, and not during the warmest period of the year as previously suggested. The achievements of photoperiod could act as a growth constraint or a limit after which the rate of tree-ring formation tends to decrease, thus allowing plants to safely complete secondary cell wall lignification before winter.

Observations on the net carbon exchange of forests in the European Mediterranean region, measured recently by the eddy covariance method, have revived interest in a phenomenon first characterized on agricultural and forest soils in East Africa in the 1950s and 1960s by H. F. Birch and now often referred to as the "Birch effect." When soils become dry during summer because of lack of rain, as is common in regions with Mediterranean climate, or are dried in the laboratory in controlled conditions, and are then rewetted by precipitation or irrigation, there is a burst of decomposition, mineralization and release of inorganic nitrogen and CO(2). In forests in Mediterranean climates in southern Europe, this effect has been observed with eddy covariance techniques and soil respiration chambers at the stand and small plot scales, respectively. Following the early work of Birch, laboratory incubations of soils at controlled temperatures and water contents have been used to characterize CO(2) release following the rewetting of dry soils. A simple empirical model based on laboratory incubations demonstrates that the amount of carbon mineralized over one year can be predicted from soil temperature and precipitation regime, provided that carbon lost as CO(2) is taken into account. We show that the amount of carbon returned to the atmosphere following soil rewetting can reduce significantly the annual net carbon gain by Mediterranean forests.

We studied the effects of three nitrogen (N) supply rates (low, intermediate and high) on Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings and poplar clone "I-214" (Populus x euroamericana (Dole) Guinier) cuttings growing in mini-stands. Our specific objectives were to: (1) evaluate the effects of N supply on water-use efficiency (WUE) and biomass production; (2) determine if N affects WUE through control of carbon assimilation rates or through stomatal control of water loss; and (3) compare three methods of estimating WUE: one short-term method (WUE(i), based on gas exchange measurements) and two long-term methods (WUE(T), based on the ratio between biomass production and transpired water, and Delta, based on leaf carbon isotope discrimination tested as a proxy of WUE). In both species, biomass production, WUE(i) and WUE(T) increased with increasing N supply, but there was no effect of N supply on either transpiration or stomatal conductance and Delta was negatively related to leaf N concentration. Plots of Delta versus both WUE(i) and WUE(T) revealed negative trends, but the regression between WUE(i) and Delta was significant only for Douglas-fir, and the regression between WUE(T) and Delta was significant only for poplar. Thus, the mechanisms underlying the response of WUE to N supply were mainly related to a positive effect of N supply on photosynthetic rates. The data confirm that carbon isotope discrimination may be a useful proxy of WUE. The finding that N availability enhances both biomass production and WUE may have practical implications in regions where these factors impose constraints on forest productivity.

The effects of harvest on European forest net ecosystem exchange (NEE) of carbon and its photosynthetic and respiratory components (GPP (gross primary production) and TER (total ecosystem respiration)) were examined by comparing four pairs of mature/harvested sites in Europe via a combination of eddy covariance measurements and empirical modeling. Three of the comparisons represented high coniferous forestry (spruce in Britain, and pines in Finland and France), while a coppice-with-standard oak plantation was examined in Italy. While every comparison revealed that harvesting converted a mature forest carbon sink into a carbon source of similar magnitude, the mechanisms by which this occurred were very different according to species or management practice. In Britain, Finland, and France the annual sink (source) strength for mature (clear-cut) stands was estimated at 496 (112), 138 (239), and 222 (225) g C m−2, respectively, with 381 (427) g C m−2 for the mature (coppiced) stand in Italy. In all three cases of high forestry in Britain, Finland, and France, clear-cutting crippled the photosynthetic capacity of the ecosystem – with mature (clear-cut) GPP of 1970 (988), 1010 (363), and 1600 (602) g C m−2– and also reduced ecosystem respiration to a lesser degree – TER of 1385 (1100), 839 (603), and 1415 (878) g C m−2, respectively. By contrast, harvesting of the coppice oak system provoked a burst in respiration – with mature (clear-cut) TER estimated at 1160 (2220) gC m−2– which endured for the 3 years sampled postharvest. The harvest disturbance also reduced GPP in the coppice system – with mature (clear-cut) GPP of 1600 (1420) g C m−2– but to a lesser extent than in the coniferous forests, and with near-complete recovery within a few years. Understanding the effects of harvest on the carbon balance of European forest systems is a necessary step towards characterizing carbon exchange for timberlands on large scales

To assess the variation of soil respiration at different forest stages we measured it in a coppiced oak (Quercus cerris L.) chronosequence in central Italy during two campaigns, spanning 2 successive years, in four stands at different stages of the rotation: 1 year (S1), 5 years (S5), 10 years (S10) and 17 years (S17) after coppicing. The contribution of the different components of soil respiration flux (aboveground litter, belowground decomposition soil organic matter and root respiration) was estimated by a paired comparison of manipulative experiments between the recently coppiced stand (S1) and mature stand (S17).Ninety percent of soil respiration values were between 1.7 and 7.8 lmol m À2 s À1 , with an overall mean ( AE SD) of 4.0 AE 2.7 lmol m À2 s À1 . Spatial variation of soil respiration was high (CV 5 44.9%), with a mean range (i.e. patch size) of 4.8 AE 2.7 m, as estimated from a semivariance analysis.In the absence of limitation by soil moisture, soil respiration was related to soil temperature with the exponential Q 10 model (average Q 10 5 2.25). During summer, soil moisture constrained soil respiration and masked its dependence on soil temperature. Soil respiration declined over the years after coppicing. Assuming a linear decline with stand age, we estimated a reduction of 24% over a 20-year-rotation cycle. The response of soil respiration to temperature also changed with age of the stands: the Q 10 was estimated to decrease from 2.90 in S1 to 2.42 in S17, suggesting that different components or processes may be involved at different developmental stages. The contribution of heterotrophic respiration to total soil respiration flux was relatively larger in the young S1 stand than in the mature S17 stand.

We compared photosynthesis-nitrogen relationships of one broad-leaved (poplar; Populus x euroamericana (Dole) Guinier) and one conifer (Douglas-fir; Pseudotsuga menziesii (Mirb.) Franco) species. Plants were grown in large pots to allow free root development and were kept well watered. We determined effects of low, intermediate and high nitrogen supply rates on area-based leaf nitrogen (Na) and chlorophyll concentrations, leaf mass per area (LMA), light-saturated photosynthesis (Amax), maximum carboxylation (Vcmax) and electron transport rate (Jmax), photosynthetic nitrogen-use efficiency (PNUE), and proportions of leaf N in active Rubisco (PR), bioenergetic pools (PB) and the light-harvesting complex (PLH). Nitrogen supply significantly affected leaf Na. Leaf mass per area did not differ between species and was unaffected by the N treatments. In both species, there was a positive correlation between leaf Na and chlorophyll concentration, and between leaf Na and the photosynthetic parameters Amax, Jmax and Vcmax. At comparable leaf Na, however, poplar showed twofold higher PNUE and a threefold steeper slope of the Amax- nitrogen relationship than Douglas-fir. Leaf Na was negatively correlated with PNUE in Douglas-fir but not in poplar. Leaf Na was also negatively correlated with PR, PB and PLH in Douglas-fir, whereas in poplar, a negative correlation was found only for PLH. Parameter PR was significantly higher in poplar than in Douglas-fir. The ratio of CO2 concentration in the intercellular space to that in ambient air was higher in poplar than in Douglas-fir. Overall, our data suggest that differences in the photosynthesis-nitrogen relationship and PNUE between Douglas-fir and poplar primarily reflect a different investment of N to active Rubisco, and possibly a different constraint to CO2 diffusion.

The present study was carried out to elucidate the response mechanisms of 50-year-old Pinus halepensis Mill. trees to a long-term and severe drought. The amount of water available to trees was artificially restricted for 12 months by covering the soil with a plastic roof. Over the short term a direct and rapid impact of drought was evident on the water relations and gas exchanges of trees: as the soil dried out in the Spring, there was a concurrent decrease of predawn water potential; transpiration was strongly reduced by stomatal closure. Seasonal changes in the water volume fractions of twig and stem xylem were observed and interpreted as the result of cavitation and refilling in the xylem. When droughted trees recovered to a more favourable water status, refilling of embolized xylem was observed; twig predawn water potentials were still negative in the period when the embolism was reversed in the twig xylem. A few months after the removal of the covering, no differences in whole plant hydraulic resistance were observed between droughted and control trees. Needle and shoot elongation and stem radial growth were considerably reduced in droughted trees; no strategy of trees to allocate carbon preferentially to the stem conducting tissues was apparent throughout the experiment. An after-effect of the drought on growth was observed