Search

Oxidative respiration is strongly temperature driven. However, in woody stems, efflux of CO(2) to the atmosphere (E (A)), commonly used to estimate the rate of respiration (R (S)), and stem temperature (T (st)) have often been poorly correlated, which we hypothesized was due to transport of respired CO(2) in xylem sap, especially under high rates of sap flow (f (s)). To test this, we measured E (A), T (st), f (s) and xylem sap CO(2) concentrations ([CO(2)*]) in 3-year-old Populus deltoides trees under different weather conditions (sunny and rainy days) in autumn. We also calculated R (S) by mass balance as the sum of both outward and internal CO(2) fluxes and hypothesized that R (S) would correlate better with T (st) than E (A). We found that E (A) sometimes correlated well with T (st), but not on sunny mornings and afternoons or on rainy days. When the temperature effect on E (A) was accounted for, a clear positive relationship between E (A) and xylem [CO(2)*] was found. [CO(2)*] varied diurnally and increased substantially at night and during periods of rain. Changes in [CO(2)*] were related to changes in f (s) but not T (st). We conclude that changes in both respiration and internal CO(2) transport altered E (A). The dominant component flux of R (S) was E (A). However, on a 24-h basis, the internal transport flux represented 9-18% and 3-7% of R (S) on sunny and rainy days, respectively, indicating that the contribution of stem respiration to forest C balance may be larger than previously estimated based on E (A) measurements. Unexpectedly, the relationship between R (S) and T (st) was sometimes weak in two of the three trees. We conclude that in addition to temperature, other factors such as water deficits or substrate availability exert control on the rate of stem respiration so that simple temperature functions are not sufficient to predict stem respiration.

We estimated daily use of stored water by Scots pine (Pinus sylvestris L.) trees growing in a temperate climate with the ANAFORE model (ANAlysis of FORest Ecosystems) and compared the simulation results with sap flow measurements. The original model was expanded with a dynamic water flow and storage model that simulates sap flow dynamics in an individual tree. ANAFORE was able to accurately simulate diurnal patterns of measured sap flow under microclimatic conditions that differ from those of the calibration period. Strong relationships were found between stored water use and several tree characteristics (diameter at breast height, sapwood area, leaf area), but not with tree height. Relative to transpiration, stored water use varied over time (between < 1% and 44% of daily transpiration). On days when transpiration was high, trees were more dependent on stored water, indicating that the contribution of internal water to transpiration is not a constant in the water budget of trees.

Rates of CO 2 efflux of stems and branches are highly variable among and within trees and across stands. Scaling factors have only partially succeeded in accounting for the observed variations. In this study, the resistance to radial CO 2 diffusion was quantified for tree stems of an eastern cottonwood (Populus deltoides Bartr. ex Marsh.) clone by direct manipulation of the CO 2 concentration ([CO 2 ]) of xylem sap under controlled conditions. Tree-specific linear relationships between rates of stem CO 2 efflux (J O ) and xylem [CO 2 ] were found. The resistance to radial CO 2 diffusion differed 6-fold among the trees and influenced the balance between the amount of CO 2 retained in the xylem v. that which diffused to the atmosphere. Therefore, we hypothesised that variability in the resistance to radial CO 2 diffusion might be an overlooked cause for the inconsistencies and large variations in woody tissue CO 2 efflux. It was found that transition from light to dark conditions caused a rapid increase in J O and xylem [CO 2 ], both in manipulated trees and in an intact tree with no sap manipulation. This resulted in an increased resistance to radial CO 2 diffusion during the dark, at least for trees with smaller daytime resistances. Stem diameter changes measured in the intact tree supported the idea that higher actual respiration rates occurred at night owing to higher metabolism in relation to an improved water status and higher turgor pressure.

Soil CO2 efflux (SR) is the second largest carbon flux on earth. We investigated the driving factors of the seasonal change and short-distance spatial variation in SR in a short-rotation plantation of willow (Salix viminalis Orm). Total annual SR ranged from 723 to 1149 g Cm(-2) year(-1). Both an exponential and a logistic model were fitted to the data, with soil temperature at a depth of 5 cm as the independent variable. The R2 values for individual sampling points ranged from 0.83 to 0.95 and from 0.85 to 0.93 for the exponential and logistic models, respectively, indicating that soil temperature largely determined the seasonal variation in SR. Modeled soil SR at 10 degrees C ranged from 1.22 to 1.95 micromol m(-2) s(-1), whereas modeled annual Q(10) values were between 3.31 and 6.13. These high Q(10) values were attributed to the absence of drought during the study in 2005. When the coefficients of the general SR models were replaced by linear dependencies on soil and vegetation-related characteristics, the resulting spatially explicit exponential and logistic SR models explained 85 and 86%, respectively, of the variability within the dataset. The analysis indicated that soil carbon concentration, leaf area index, soil pH and root biomass caused differences in SR at the short distances considered in this study. However, incorporating information on variables considered to account for spatial variability in the model did not result in a higher R2 compared with a simple temperature function. When the general SR models were applied to independent datasets from the same plantation, the logistic model provided a better fit than the exponential model when drought occurred. Drought greatly reduced the annual Q(10) values of SR.

A young potted oak (Quercus robur L.) tree was subjected to drought by interrupting the water supply for 9 days. The tree was placed in a growth chamber in which daily patterns of temperature and radiation were constant. The effects of drought on the water and carbon status of the stem were examined by measuring stem sap flow rate, stem water potential, stem diameter variations, stem CO(2) efflux rate (F(CO2)) and xylem CO(2) concentration ([CO(2)*]). Before and after the drought treatment, diurnal fluctuations in F(CO2) and [CO(2)*] corresponded well with variations in stem temperature (T(st)). Daytime depressions in F(CO2) did not occur. During the drought treatment, F(CO2) still responded to stepwise changes in temperature, but diurnal fluctuations in F(CO2) were no longer correlated with diurnal fluctuations in T(st). From the moment daily growth rate of the stem became zero, diurnal fluctuations in F(CO2) became closely correlated with diameter variations, exhibiting clear daytime depressions. The depressions in F(CO2) were likely the result of a reduction in metabolic activity caused by the lowered daytime stem water status. Xylem [CO(2)*] showed clear daytime depressions in response to drought. When the tree was re-watered, F(CO2) and [CO(2)*] exhibited sharp increases, coinciding with an increase in stem diameter. After resumption of the water supply, daytime depressions in F(CO2) and [CO(2)*] disappeared and diurnal fluctuations in F(CO2) and [CO(2)*] corresponded again with variations in T(st).

Background and Aims: Daytime CO2 efflux rates (FCO2) from tree stems are often reported to be lower than expected from the exponential relationship between temperature and respiration. Explanations of daytime depression in FCO2 have focused on the possible role of internal CO2 transport in the xylem. However, another possible cause that has been overlooked is the daily dynamics of the water status in the living stem tissues and its influence on stem growth rate and thus respiration. The objective of this study was to assess the daily dynamics of stem water status and growth rate and to determine the extent to which they may be linked to daily variations in stem FCO2., Methods: FCO2 of young beech and oak stems were measured under controlled conditions. Relative stem turgor pressure (Psi(p)), obtained from simulations with the 'RCGro' model, was used as an indicator of the water status in the living stem tissues. Daily dynamics of stem growth were derived from Psi(p): growth was assumed to occur when Psi(p) exceeded a relative threshold value., Key Results: There was a strong correspondence between fluctuations in FCO2 and simulated Psi(p). The non-growth conditions during daytime coincided with depressions in FCO2. Moreover, FCO2 responded to changes in Psi(p) in the absence of growth, indicating also that maintenance processes were influenced by the water status in the living stem tissues., Conclusions: Daytime depressions in stem FCO2 correlate with the daily dynamics of turgor, as a measure of the water status in the living stem tissues: it is suggested that water status of tree stems is a potentially important determinant of stem FCO2, as it influences the rate of growth and maintenance processes in the living tissues of the stem.

Calibration of a recently developed water flow and storage model based on experimental data for a young diffuse-porous beech tree (Fagus sylvatica L.) and a young ring-porous oak tree (Quercus robur L.) revealed that differences in stem wood anatomy between species strongly affect the calibrated values of the hydraulic model parameters. The hydraulic capacitance (C) of the stem storage tissue was higher in oak than in beech (939.8 versus 212.3 mg MPa(-1)). Model simulation of the elastic modulus (epsilon) revealed that this difference was linked to the higher elasticity of the stem storage tissue of oak compared with beech. Furthermore, the hydraulic resistance (R (x)) of beech was about twice that of oak (0.1829 versus 0.1072 MPa s mg(-1)). To determine the physiological meaning of the R (x) parameter identified by model calibration, we analyzed the stem wood anatomy of the beech and oak trees. Calculation of stem specific hydraulic conductivity (k (s)) of beech and oak with the Hagen-Poiseuille equation confirmed the differences in R (x) predicted by the model. The contributions of different vessel diameter classes to the total hydraulic conductivity of the xylem were calculated. As expected, the few big vessels contributed much more to total conductivity than the many small vessels. Compared with beech, the larger vessels of oak resulted in a higher k (s) (10.66 versus 4.90 kg m(-1) s(-1) MPa(-1)). The calculated ratio of k (s) of oak to beech was 2, confirming the R (x) ratio obtained by model calibration. Thus, validation of the R (x) parameter of the model led to identification of its physiological meaning.

The Monte Carlo technique can be used to propagate input variable uncertainty and parameter uncertainty through a model to determine output uncertainty. However, to carry out Monte Carlo simulations, the uncertainty distributions or the probability density functions (PDFs) of the model parameters and input variables must be known. This remains one of the bottlenecks in current uncertainty research in forest carbon flux modeling. Because forest carbon flux models involve many parameters, we questioned whether it is necessary to take into account all parameters in the uncertainty analysis. A sensitivity analysis can determine the parameters contributing most to the overall model output uncertainty. This paper illustrates the usefulness of the Monte Carlo simulation technique for ranking parameters for sensitivity and uncertainty in process-based forest flux models. The uncertainty of the output (net ecosystem exchange, NEE) of the FORUG model was estimated for the Hesse beech forest (1997). Based on the arbitrary uncertainty of ten key parameters, a standard deviation of 0.88 Mg C ha(-1) year(-1) NEE was found which is equal to 24% of the mean value of NEE. Sensitivity analysis showed that the overall output uncertainty of the FORUG model can largely be determined by accounting for the uncertainty of only a few key parameters. The results led to the identification of the key FORUG parameters and to the recommendation for a process-based description of the soil respiration process in the FORUG model.

Background and Aims: Stomatal oscillations have been reported in many plant species, but they are usually induced by sudden step changes in the environment when plants are grown under constant conditions. This study shows that in navel orange trees (Citrus sinensis) pronounced stomatal oscillations occur and persist under natural climatic conditions., Methods: Oscillations in stomatal conductance were measured, and related to simultaneous measurements of leaf water potential, and flow rate of sap in the stems of young, potted plants. Cycling was also observed in soil-grown, mature orchard trees, as indicated by sap flow in stem and branches., Key Results: Oscillations in stomatal conductance were caused by the rapid propagation and synchronization of changes in xylem water potential throughout the tree, without rapid changes in atmospheric conditions., Conclusions: The results show marked stomatal oscillations persisting under natural climatic conditions and underscore the need to discover why this phenomenon is so pronounced in orange trees.

To date, models for simulating sap flow dynamics in individual trees with a direct link to stem diameter variation include only the diameter fluctuation driven by a change in stem water storage. This paper reports results obtained with a comprehensive flow and storage model using whole-tree leaf transpiration as the only input variable. The model includes radial stem growth based on Lockhart's equation for irreversible cell expansion. It was demonstrated that including growth is essential to obtaining good simulation results. To model sap flow dynamics, capacitance of storage tissues was assumed either constant (i.e., electrical analogue approach) or variable and dependent on the water content of the respective storage tissue (i.e., hydraulic system approach). These approaches resulted in different shapes for the desorption curve used to calculate the capacitance of storage tissues. Comparison of these methods allowed detection of specific differences in model simulation of sap flow at the stem base (F(stem)) and stem diameter variation (D). Sensitivity analysis was performed to select a limited subset of identifiable parameters driving most of the variability in model predictions of F(stem) and D Both the electrical analogue and the hydraulic system approach for the flow and storage model were successfully calibrated and validated for the case of a young beech tree (Fagus sylvatica L.). Use of an objective model selection criterion revealed that the flow and storage model based on the electrical analogue approach yielded better predictions.

Results from measurement of sap flow by heat balance sensors on the stem of a young oak tree (Quercus robur L.) revealed that thermal disequilibrium (i.e. heat storage) within the heated stem segment can introduce considerable errors in the measured sap-flow rates. The magnitude, sign and significance of these errors depend on the sap-flow rate and on the relationship between stem temperature and air temperature. Sap-flow rates were found to be more prone to errors caused by heat storage effects under low flow conditions than at higher rates of sap flow. Furthermore, daytime fluctuations of air temperature and stem temperature inside the heat balance sensor were either in phase when a low, or in opposite phase when a high sap-flow rate was passing through the stem of the young tree. To investigate this relationship, we developed an experimental set-up with cut stem segments through which tap water could be pressed. This set-up allowed the effects of air temperature and sap-flow rates on stem temperatures within heat balance sensors to be clearly separated. Good mathematical relationships were obtained and were successfully used to assess the relative importance of air temperature and sap-flow rate with respect to the fluctuations in stem temperature of the young oak tree. Based on the established relationships, a novel approach was put forward to correct for errors introduced into sap-flow calculations caused by heat storage effects if no measured data on stem temperature are available.

Quantitative analysis of wood anatomical characteristics is usually performed using classical microtomy yielding optical micrographs of stained thin sections. It is time-consuming to obtain high quality cross-sections from microtomy, and sections can be damaged. This approach, therefore, is often impractical for those who need quick acquisition of quantitative data on vessel characteristics in wood. This paper reports results of a novel approach using X-ray computed microtomography (microCT) for non-invasive determination of wood anatomy. As a case study, stem wood samples of a 2-year-old beech (Fagus sylvatica L.) and a 3-year-old oak (Quercus robur L.) tree were investigated with this technique, beech being a diffuse-porous and oak a ring-porous tree species. MicroCT allowed non-invasive mapping of 2-D transverse cross-sections of both wood samples with micrometer resolution. Self-developed software 'microCTanalysis' was used for image processing of the 2-D cross-sections in order to automatically determine the inner vessel diameters, the transverse cross-sectional surface area of the vessels, the vessel density and the porosity with computer assistance. Performance of this new software was compared with manual analysis of the same micrographs. The automatically obtained results showed no significant statistical differences compared to the manual measurements. Visual inspection of the microCT slices revealed very good correspondence with the optical micrographs. Statistical analysis confirmed this observation in a more quantitative way, and it was, therefore, argued that anatomical analysis of optical micrographs can be readily substituted by automated use of microCT, and this without loss of accuracy. Furthermore, as an additional application of microCT, the 3-D renderings of the internal microstructure of the xylem vessels for both the beech and the oak sample could be reconstructed, clearly showing the complex nature of vessel networks. It can be concluded that the use of microCT in wood science offers an interesting potential for all those who need quantitative data of wood anatomical characteristics in either the 2-D or the 3-D space.

This paper describes an experimental system designed for analysis of the dynamic water flow through young trees, complemented with test results of a 2-year-old beech tree (Fagus sylvatica L.). The system allows automatic and simultaneous measurements of a complex set of plant physiological processes at leaf, branch, stem and root levels [transpiration (E), sap flow (F) and diameter fluctuations (Δd)], in combination with the micrometeorological variables that control these processes [soil and air temperature (T s and T a ), vapour pressure deficit of the air (D) and photosynthetically active radiation (PAR)]. A 2-d experimental period was used to study the whole-tree water transport dynamics of the young beech tree. Good correspondence between E of the leaves and F in the supporting branch was found. An increased time lag between the F measurements along the hydraulic pathway down towards the root system was observed, indicating the non-steady-state nature of the water flow. The daytime Δd of stem and branch revealed the depletion and the replenishment of internal water reserves. The daily amount of water withdrawn from internal storage was 5% of the total daily transpiration. A good linear relationship was found between the rate of change in internal water storage and the rate of change in stem diameter, having no time lag. We conclude that the data obtained with this system will allow experimental assessment of hydraulic properties in young trees and facilitate calibration of models for non-steady-state conditions of water flow in young trees.