Fertilization effects on mean stomatal conductance are mediated through changes in the hydraulic attributes of mature Norway spruce trees.

Stomatal conductance was quantified with sap flux sensors and whole-tree chambers in mature Norway spruce (Picea abies (L.) Karst.) trees after 3 years of exposure to elevated CO(2) concentration ([CO(2)]) in a 13-year nutrient optimization experiment. The long-term nutrient optimization treatment increased tree height by 3.7 m (67%) and basal diameter by 8 cm (68%); the short-term elevated [CO(2)] exposure had no effect on tree size or allometry. Nighttime transpiration was estimated as approximately 7% of daily transpiration in unchambered trees; accounting for the effect of nighttime flux on the processing of sap flux signals increased estimated daily water uptake by approximately 30%. Crown averaged stomatal conductance (g(s)) was described by a Jarvis-type model. The addition of a stomatal response time constant (tau) and total capacitance of stored water (C(tot)) improved the fit of the model. Model estimates for C(tot) scaled with sapwood volume of the bole in fertilized trees. Hydraulic support-defined as a lumped variable of leaf-specific hydraulic conductivity and water potential gradient (K(l)DeltaPsi) -was estimated from height, sapwood-to-leaf area ratio (A(s):A(l)) and changes in tracheid dimensions. Hydraulic support explained 55% of the variation in g(s) at reference conditions for trees across nutrient and [CO(2)] treatments. Removal of approximately 50% of A(l) from three trees yielded results suggesting that stomatal compensation (i.e., an increase in g(s)) after pruning scales inversely with K(l)DeltaPsi, indicating that the higher the potential hydraulic support after pruning, the less complete the stomatal compensation for the increase in A(s):A(l).

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 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).

Variations in 13C discrimination during CO2 exchange by Picea sitchensis branches in the field.

We report diurnal variations in (13)C discrimination ((13)Delta) of Picea sitchensis (Bong.) Carr. branches measured in the field using a branch chamber technique. The observations were compared to predicted (13)Delta based on concurrent measurements of branch gas exchange. Observed (13)Delta values were described well by the classical model of (13)Delta including isotope effects during photorespiration, day respiration and CO(2) transfer through a series of resistances to the sites of carboxylation. A simplified linear of model (13)Delta did not capture the observed diurnal variability. At dawn and dusk, we measured very high (13)Delta values that were not predicted by either of the said models. Exploring the sensitivity of (13)Delta to possible respiratory isotope effects, we conclude that isotopic disequilibria between the gross fluxes of photosynthesis and day respiration can explain the high observed (13)Delta values during net photosynthetic gas exchange. Based on the classical model, a revised formulation incorporating an isotopically distinct substrate for day respiration was able to account well for the high observed dawn and dusk (13)Delta values.

The likely impact of elevated [CO2], nitrogen deposition, increased temperature and management on carbon sequestration in temperate and boreal forest ecosystems: a literature review.

Temperate and boreal forest ecosystems contain a large part of the carbon stored on land, in the form of both biomass and soil organic matter. Increasing atmospheric [CO2], increasing temperature, elevated nitrogen deposition and intensified management will change this C store. Well documented single-factor responses of net primary production are: higher photosynthetic rate (the main [CO2] response); increasing length of growing season (the main temperature response); and higher leaf-area index (the main N deposition and partly [CO2] response). Soil organic matter will increase with increasing litter input, although priming may decrease the soil C stock initially, but litter quality effects should be minimal (response to [CO2], N deposition, and temperature); will decrease because of increasing temperature; and will increase because of retardation of decomposition with N deposition, although the rate of decomposition of high-quality litter can be increased and that of low-quality litter decreased. Single-factor responses can be misleading because of interactions between factors, in particular those between N and other factors, and indirect effects such as increased N availability from temperature-induced decomposition. In the long term the strength of feedbacks, for example the increasing demand for N from increased growth, will dominate over short-term responses to single factors. However, management has considerable potential for controlling the C store.

A comparative analysis of simulated and observed photosynthetic CO2 uptake in two coniferous forest canopies.

Gross canopy photosynthesis (P(g)) can be simulated with canopy models or retrieved from turbulent carbon dioxide (CO2) flux measurements above the forest canopy. We compare the two estimates and illustrate our findings with two case studies. We used the three-dimensional canopy model MAESTRA to simulate P(g) of two spruce forests differing in age and structure. Model parameter acquisition and model sensitivity to selected model parameters are described, and modeled results are compared with independent flux estimates. Despite higher photon fluxes at the site, an older German Norway spruce (Picea abies L. (Karst.)) canopy took up 25% less CO2 from the atmosphere than a young Scottish Sitka spruce (Picea sitchensis (Bong.) Carr.) plantation. The average magnitudes of P(g) and the differences between the two canopies were satisfactorily represented by the model. The main reasons for the different uptake rates were a slightly smaller quantum yield and lower absorptance of the Norway spruce stand because of a more clumped canopy structure. The model did not represent the scatter in the turbulent CO2 flux densities, which was of the same order of magnitude as the non-photosynthetically-active-radiation-induced biophysical variability in the simulated P(g). Analysis of residuals identified only small systematic differences between the modeled flux estimates and turbulent flux measurements at high vapor pressure saturation deficits. The merits and limitations of comparative analysis for quality evaluation of both methods are discussed. From this analysis, we recommend use of both parameter sets and model structure as a basis for future applications and model development.

Measurement of CO2 exchange between Boreal forest and the atmosphere.

The Boreal forest is the world's second largest forested biome occupying the circumpolar region between 50 degrees N and 70 degrees N. This heterogeneous biome stores about 25% of all terrestrial carbon. We have reviewed EC measurements of CO2 exchange between the atmosphere and Boreal forests, and assessed progress in understanding the controlling processes. We have assessed net ecosystem productivity, the net balance between net primary productivity and heterotrophic respiration, measured using the EC method, for 38 Boreal forest sites. Gross ecosystem productivity has been estimated by adding day-time EC-measured CO2 fluxes to respiration estimated from night-time relationships between respiration and temperature. Maximum midday values of gross ecosystem productivity vary from 33 pmol m(-2) s(-1) for aspen to 6 micromol m(-2) s(-1) for larch stands. Long-term EC flux measurements, ongoing at nine Boreal sites, have shown the strong impact of spring weather and growing season water balance on annual net ecosystem productivity. Estimation of net biome production, incorporating the effects of disturbance resulting from forest fires and logging, has progressed significantly in recent years. After disturbance, summer measurements in Boreal chronosequences suggest that it takes about 10 years before growing season carbon uptake offsets the decomposition emissions. Small-scale exchange rate measurements using chambers and manipulative experiments such as stem girdling and soil heating help to understand the processes and mechanisms playing major roles in the carbon balance of terrestrial ecosystems. Aircraft EC flux measurements, convective boundary layer carbon budgets, and (13)C/12C changes in the atmosphere play an important role in validating estimates of regional carbon exchange based on scaled up EC measurements. Atmospheric inverse models are an important approach to studying regional and global carbon balance but need further improvement to yield reliable quantitative results.

The carbon balance of forest soils: detectability of changes in soil carbon stocks in temperate and Boreal forests.

Estimating soil carbon content as the product of mean carbon concentration and bulk density can result in considerable overestimation. Carbon concentration and soil mass need to be measured on the same sample and carbon contents calculated for each individual sample before averaging. The effect of this bias is likely to be smaller (but still greater than zero) when the primary objective is to determine stock changes over time. Variance and mean carbon content are significantly and positively related to each other, although some sites showed much higher variability than predicted by this relationship, as a likely consequence of their particular site history, forest management, and micro-topography. Because of the proportionality between mean and variance, the number of samples required to detect a fixed change in soil carbon stocks varied directly with the site mean carbon content from less than 10 to several thousands across the range of carbon stocks normally encountered in temperate and Boreal forests. This raises important questions about how to derive an optimal sampling strategy across such a varied range of conditions so as to achieve the aims of the Kyoto Protocol. Overall, on carbon-poor forest sites with little or no disturbance to the soil profile, it is possible to detect changes in total soil organic carbon over time of the order of 0.5 kg (C) m(-2) with manageable sample sizes even using simple random sampling (i.e., about 50 samples per sampling point). More efficient strategies will reveal even smaller differences. On disturbed forest sites (ploughed, windthrow) this is no longer possible (required sample sizes are much larger than 100). Soils developed on coarse aeolian sediments (sand dunes), or where buried logs or harvest residues of the previous rotation are present, can also exhibit large spatial variability in soil carbon. Generally, carbon-rich soils will always require larger numbers of samples. On these sites, simple random sampling is unlikely to be the preferred method, because of its inherent inefficiency. More sophisticated approaches, such as paired re-sampling inside relatively small plots (see, for example, Ellert et al., 2001) are likely to reduce sample size significantly and lead to detection of smaller differences in carbon stocks over time. However, it remains to be shown that at these sites the application of efficient sampling designs will result in the detection of differences relevant for the objectives of the Kyoto Protocol (cf., Conant et al., 2003). Finally, it should also be noted that, compared to the accuracy with which changes in atmospheric carbon content can be detected (less than 1 p.p.m. CO2), changes in soil carbon stocks are very uncertain. A release of 0.5 kg (C) from 1 m2 of soil surface is equivalent to an increase in CO, concentration of about 125 p.p.m. in the air column above the same area.

Gas exchange, biomass, whole-plant water-use efficiency and water uptake of peach (Prunus persica) seedlings in response to elevated carbon dioxide concentration and water availability.

We examined the interactive effects of elevated CO2 concentration ([CO2]) and water stress on growth and physiology of 1-year-old peach (Prunus persica L.) seedlings grown in 10-dm3 pots in open-top chambers with ambient (350 micromol mol-1) or elevated (700 micromol mol-1) [CO2]. Seedlings were supplied weekly with a non-limiting nutrient solution. Water was withheld from half of the plants in each treatment for a 4-week drying cycle, to simulate a sudden and severe water stress during the phase of rapid plant growth. Throughout the growing season, seedlings in elevated [CO2] had higher assimilation rates, measured at the growth [CO2], than seedlings in ambient [CO2], and this caused an increase in total dry mass of about 33%. Stomatal conductance, total water uptake, leaf area and leaf number were unaffected by elevated [CO2]. Because seedlings in the two CO2 treatments had similar transpiration despite large differences in total dry mass, water-use efficiency (WUE) of well-watered and water-stressed seedlings grown in elevated [CO2] was an average of 51 and 63% higher, respectively, than WUE of comparable seedlings grown in ambient [CO2]. Elevated [CO2] enhanced total biomass of water-stressed seedlings by 31%, and thus ameliorated the effects of water limitation. However, the percentage increases in total dry mass between well-watered and water-stressed seedlings were similar in ambient (53%) and elevated (58%) [CO2], demonstrating that there was no interaction between elevated [CO2] and water stress. This finding should be considered when predicting responses of trees to global climate change in hot and dry environments, where predicted temperature increases will raise evaporative demands and exacerbate the effects of drought on tree growth.

Influence of male and female cones on assimilate production of Pinus contorta trees within a forest stand.

Studies on branches of field-grown Pinus contorta Dougl. trees showed that: (i) branches with fully developed male or female cones were heavier than vegetative branches; (ii) the production of female cones decreased neither the number of laterals nor the length of the terminal or lateral shoots on the cone-bearing branches; (iii) the production of male cones decreased both the number of laterals and the number of needles on the cone-bearing branches; (iv) needles on male cone-bearing branches had significantly higher photosynthetic rates and needles on female cone-bearing branches generally also fixed more CO(2) per quantum of light. In addition, both male and female cones refixed a significant proportion of respired CO(2) when illuminated. The simulation model, MAESTRO, was used to estimate light penetration through two stands (3690 and 1845 trees ha(-1)) of P. contorta trees and to calculate the resultant CO(2) assimilation of a vegetative, a male and a female cone-bearing tree on sunny, partly sunny and cloudy days in Scotland. Generally, the CO(2) efflux from female cones was less than 3% of the CO(2) assimilated by the whole tree. The presence of male cones resulted in an average 33% decrease in needle complement, but an increased quantum efficiency of associated needles in the autumn. It was estimated that, on a sunny day, male cone-bearing trees assimilated as much CO(2) as vegetative trees in the high density stand and only 6% less in the low density stand. On cloudy days, the smaller needle complement was beneficial to the carbon economy of the tree because of lower respiration losses compared with a vegetative tree.