Canopy nitrogen, carbon assimilation, and albedo in temperate and boreal forests: Functional relations and potential climate feedbacks.

The availability of nitrogen represents a key constraint on carbon cycling in terrestrial ecosystems, and it is largely in this capacity that the role of N in the Earth's climate system has been considered. Despite this, few studies have included continuous variation in plant N status as a driver of broad-scale carbon cycle analyses. This is partly because of uncertainties in how leaf-level physiological relationships scale to whole ecosystems and because methods for regional to continental detection of plant N concentrations have yet to be developed. Here, we show that ecosystem CO(2) uptake capacity in temperate and boreal forests scales directly with whole-canopy N concentrations, mirroring a leaf-level trend that has been observed for woody plants worldwide. We further show that both CO(2) uptake capacity and canopy N concentration are strongly and positively correlated with shortwave surface albedo. These results suggest that N plays an additional, and overlooked, role in the climate system via its influence on vegetation reflectivity and shortwave surface energy exchange. We also demonstrate that much of the spatial variation in canopy N can be detected by using broad-band satellite sensors, offering a means through which these findings can be applied toward improved application of coupled carbon cycle-climate models.

Uncertainty in eddy covariance measurements and its application to physiological models.

Flux data are noisy, and this uncertainty is largely due to random measurement error. Knowledge of uncertainty is essential for the statistical evaluation of modeled and measured fluxes, for comparison of parameters derived by fitting models to measured fluxes and in formal data-assimilation efforts. We used the difference between simultaneous measurements from two towers located less than 1 km apart to quantify the distributional characteristics of the measurement error in fluxes of carbon dioxide (CO2) and sensible and latent heat (H and LE, respectively). Flux measurement error more closely follows a double exponential than a normal distribution. The CO2 flux uncertainty is negatively correlated with mean wind speed, whereas uncertainty in H and LE is positively correlated with net radiation flux. Measurements from a single tower made 24 h apart under similar environmental conditions can also be used to characterize flux uncertainty. Uncertainty calculated by this method is somewhat higher than that derived from the two-tower approach. We demonstrate the use of flux uncertainty in maximum likelihood parameter estimates for simple physiological models of daytime net carbon exchange. We show that inferred model parameters are highly correlated, and that hypothesis testing is therefore possible only when the joint distribution of the model parameters is taken into account.

Leaf conductance and CO(2) assimilation of Larix gmelinii growing in an eastern Siberian boreal forest.

In July 1993, we measured leaf conductance, carbon dioxide (CO(2)) assimilation, and transpiration in a Larix gmelinii (Rupr.) Rupr. ex Kuzen forest in eastern Siberia. At the CO(2) concentration of ambient air, maximum values (mean of 10 highest measured values) for CO(2) assimilation, transpiration and leaf conductance for water vapor were 10.1 micro mol m(-2) s(-1), 3.9 mmol m(-2) s(-1) and 365 mmol m(-2) s(-1), respectively. The corresponding mean values, which were much lower than the maximum values, were 2.7 micro mol m(-2) s(-1), 1.0 mmol m(-2) s(-1) and 56 mmol m(-2) s(-1). The mean values were similar to those of Vaccinium species in the herb layer. The large differences between maximum and actual performance were the result of structural and physiological variations within the tree crowns and between trees that reduced maximum assimilation and leaf conductance by about 40 and 60%, respectively. Thus, maximum assimilation and conductance values averaged over the canopy were 6.1 micro mol m(-2) s(-1) and 146 mmol m(-2) s(-1), respectively. Dry air caused stomatal closure, which reduced assimilation by an additional 26%. Low irradiances in the morning and evening had a minor effect (-6%). Daily canopy transpiration was estimated to be 1.45 mm day(-1), which is higher than the value of 0.94 mm day(-1) measured by eddy covariance, but similar to the value of 1.45 mm day(-1) calculated from the energy balance and soil evaporation, and less than the value of 2.1 mm day(-1) measured by xylem flux. Daytime canopy carbon assimilation, expressed on a ground area basis, was 0.217 mol m(-2) day(-1), which is higher than the value measured by eddy flux (0.162 mol m(-2) day(-1) including soil respiration). We discuss the regulation of leaf gas exchange in Larix under the extreme climatic conditions of eastern Siberia (temperature > 35 degrees C and vapor pressure deficit > 5.0 kPa).

Optimality and nitrogen allocation in a tree canopy.

Physical and functional properties of foliage were measured at a variety of microsites in a broad-leaved Nothofagus fusca (Hook. f.) Ørst. canopy. The light climate of the foliage at these sites was monitored for 39 days in the late spring and early summer with in situ sensors. Foliage nitrogen content (N), mean leaf angle, and gas exchange characteristics were all correlated with the amount of light reaching the microsites during foliage development. Foliage N content on a leaf area basis ranged between ~1 and 2.5 g N m(-2) and was highest at the brightest sites. Light-saturated photosynthetic rates ranged between ~4 and 9 micro mol m(-2) s(-1), increasing from the darkest to brightest sites. A biochemical model of photosynthesis was fitted to foliage characteristics at the different microsites and used to integrate foliage assimilation among the sites over 39 days. The actual arrangement of foliage physiological characteristics in the observed microsites led to higher total canopy rates of net assimilation than > 99% of the combinations of observed foliage characteristics randomly assigned to the observed microsites. Additional simulations first related the maximum rates of electron transport (J(max)), ribulose bisphosphate turnover (V(c,max)), and dark respiration (R(d)) of Nothofagus fusca foliage to nitrogen content and then allowed foliage N (and consequently leaf gas exchange characteristics) to vary across the canopy. The observed N allocation pattern results in greater total canopy assimilation than uniform or > 99% of the simulations with random distributions of N among the microsites (constrained so that the total N allocated was equivalent to that observed in the microsites). However, the observed pattern of N allocation places less N in the brightest microsites and results in substantially less total assimilation than a simulated canopy in which N was allocated in an optimal manner where the N distribution is such that the partial derivative of leaf assimilation (A) with respect to leaf nitrogen content, partial differential A/ partial differential N, is constant among microsites. These results suggest that other factors such as wind or herbivory reduce the integrated assimilation of high-N foliage relatively more than lower-N foliage and that a partial differential A/ partial differential N optimality criteria based only on formulations of leaf gas exchange overestimate canopy assimilation.

Environmental regulation of xylem sap flow and total conductance of Larix gmelinii trees in eastern Siberia.

Xylem sap flow and environmental variables were measured on seven consecutive midsummer days in a 130-year-old Larix gmelinii (Rupr.) Rupr. forest located 160 km south of Yakutsk in eastern Siberia, Russia (61 degrees N, 128 degrees E, 300 m asl). The site received 20 mm of rainfall during the 4 days before measurements, and soil samples indicated that the trees were well watered. The tree canopy was sparse with a one-sided leaf area index of 1.5 and a tree density of 1760 ha(-1). On a clear day when air temperature ranged from 9 to 29 degrees C, and maximum air saturation deficit was 3.4 kPa, daily xylem sap flux (F) among 13 trees varied by an order of magnitude from 7 l day(-1) for subcanopy trees (representing 55% of trees in the forest) to 67 l day(-1) for emergent trees (representing 18% of trees in the forest). However, when based on xylem sap flux density (F'), calculated by dividing F by projected tree crown area (a surrogate for the occupied ground area), there was only a fourfold range in variability among the 13 trees, from 1.0 to 4.4 mm day(-1). The calculation of F' also eliminated systematic and large differences in F among emergent, canopy and subcanopy trees. Stand-level F', estimated by combining half-hourly linear relationships between F and stem cross-sectional area with tree size distribution data, ranged between 1.8 +/- 0.4 (standard deviation) and 2.3 +/- 0.6 mm day(-1). These stand-level F' values are about 0.6-0.7 mm day(-1) (30%) larger than daily tree canopy transpiration rates calculated from forest energy balance and understory evaporation measurements. Maximum total tree conductance for water vapor transfer (G(tmax), including canopy and aerodynamic conductances), calculated from the ratio of F' and the above-canopy air saturation deficit (D) for the eight trees with continuous data sets, was 9.9 +/- 2.8 mm s(-1). This is equivalent to a leaf-scale maximum stomatal conductance (g(smax)) of 6.1 mm s(-1), when expressed on a one-sided leaf area basis, which is comparable to the published porometer data for Larix. Diurnal variation in total tree conductance (G(t)) was related to changes in the above-canopy visible irradiance (Q) and D. A saturating upper-boundary function for the relationship between G(t) and Q was defined as G(t) = G(tmax)(Q/[Q + Q(50)]), where Q(50) = 164 +/- 85 micro mol m(-2) s(-1) when G(t) = G(tmax)/2. Accounting for Q by excluding data for Q < Q(85) when G(t) was at least 85% of G(tmax), the upper limit for the relationship between G(t) and D was determined based on the function G(t) = (a + blnD)(2), where a and b are regression coefficients. The relationship between G(t) and D was curvilinear, indicating that there was a proportional decrease in G(t) with increasing D such that F was relatively constant throughout much of the day, even when D ranged between about 2 and 4 kPa, which may be interpreted as an adaption of the species to its continental climate. However, at given values of Q and D, G(t) was generally higher in the morning than in the afternoon. The additional environmental constraints on G(t) imposed by leaf nitrogen nutrition and afternoon water stress are discussed.

Transpiration and canopy conductance in a pristine broad-leaved forest of Nothofagus: an analysis of xylem sap flow and eddy correlation measurements.

Tree transpiration was determined by xylem sap flow and eddy correlation measurements in a temperate broad-leaved forest of Nothofagus in New Zealand (tree height: up to 36 m, one-sided leaf area index: 7). Measurements were carried out on a plot which had similar stem circumference and basal area per ground area as the stand. Plot sap flux density agreed with tree canopy transpiration rate determined by the difference between above-canopy eddy correlation and forest floor lysimeter evaporation measurements. Daily sap flux varied by an order of magnitude among trees (2 to 87 kg day-1 tree-1). Over 50% of plot sap flux density originated from 3 of 14 trees which emerged 2 to 5 m above the canopy. Maximum tree transpiration rate was significantly correlated with tree height, stem sapwood area, and stem circumference. Use of water stored in the trees was minimal. It is estimated that during growth and crown development, Nothofagus allocates about 0.06 m of circumference of main tree trunk or 0.01 m2 of sapwood per kg of water transpired over one hour.Maximum total conductance for water vapour transfer (including canopy and aerodynamic conductance) of emergent trees, calculated from sap flux density and humidity measurements, was 9.5 mm s-1 that is equivalent to 112 mmol m-2 s-1 at the scale of the leaf. Artificially illuminated shoots measured in the stand with gas exchange chambers had maximum stomatal conductances of 280 mmol m-2 s-1 at the top and 150 mmol m-2 s-1 at the bottom of the canopy. The difference between canopy and leaf-level measurements is discussed with respect to effects of transpiration on humidity within the canopy. Maximum total conductance was significantly correlated with leaf nitrogen content. Mean carbon isotope ratio was -27.76±0.27‰ (average ±s.e.) indicating a moist environment. The effects of interactions between the canopy and the atmosphere on forest water use dynamics are shown by a fourfold variation in coupling of the tree canopy air saturation deficit to that of the overhead atmosphere on a typical fine day due to changes in stomatal conductance.

Gas exchange and dry matter allocation responses to elevation of atmospheric CO(2) concentration in seedlings of three tree species.

Photosynthetic rates of 13-month-old Pinus radiata D. Don, Nothofagus fusca (Hook f.) Ørst. and Pseudotsuga menziesii (Mirb.) Franco seedlings grown and measured at elevated atmospheric concentrations of CO(2) (~620 microl l(-1)) were 32 to 55% greater than those of seedlings grown and measured at ambient (~310 microl l(-1)) concentrations of CO(2). Seedlings grown in ambient and elevated concentrations of CO(2) had similar rates of photosynthesis when measured at ~620 microl l(-1) CO(2), but when measured at ~310 microl l(-1) CO(2), the P. radiata and N. fusca seedlings which were grown at elevated CO(2) had lower rates of photosynthesis than the seedlings grown at an ambient concentration of CO(2). Stomatal conductances in general were lower when measured at ~620 microl l(-1) CO(2) than at ~310 microl l(-1) CO(2). Stomatal conductances declined in all species grown at both CO(2) concentrations when the leaf-air water vapor concentration gradient (DeltaW) was increased from 10 to 20 mmol H(2)O mol(-1) air. The percent enhancement in photosynthesis for P. radiata and P. menziesii at elevated CO(2) was greater at 20 mmol than at 10 mmol DeltaW, suggesting that elevated CO(2) may moderate the effects of atmospheric water stress. Dry matter allocation patterns were not significantly different for plants grown in ambient or high CO(2) air.