Transpiration and forest structure in relation to soil waterlogging in a Hawaiian montane cloud forest

Transpiration, leaf characteristics and forest structure in Metrosideros polymorpha Gaud. stands growing in East Maui, Hawaii were investigated to assess physiological limitations associated with flooding as a mechanism of reduced canopy leaf area in waterlogged sites. Whole-tree sap flow, stomatal conductance, microclimate, soil oxidation-reduction potential, stand basal area and leaf area index (LAI) were measured on moderately sloped, drained sites with closed canopies (90%) and on level, waterlogged sites with open canopies (50-60%). The LAI was measured with a new technique based on enlarged photographs of individual tree crowns and allometric relationships. Sap flow was scaled to the stand level by multiplying basal area-normalized sap flow by stand basal area. Level sites had lower soil redox potentials, lower mean stand basal area, lower LAI, and a higher degree of soil avoidance by roots than sloped sites. Foliar nutrients and leaf mass per area (LMA) in M. polymorpha were similar between level and sloped sites. Stomatal conductance was similar for M. polymorpha saplings on both sites, but decreased with increasing tree height (r(2) = 0.72; P < 0.001). Stand transpiration estimates ranged from 79 to 89% of potential evapotranspiration (PET) for sloped sites and from 28 to 51% of PET for level sites. Stand transpiration estimates were strongly correlated with LAI (r(2) = 0.96; P < 0.001). Whole-tree transpiration was lower at level sites with waterlogged soils, but was similar or higher for trees on level sites when normalized by leaf area. Trees on level sites had a smaller leaf area per stem diameter than trees on sloped sites, suggesting that soil oxygen deficiency may reduce leaf area. However, transpiration per unit leaf area did not vary substantially, so leaf-level physiological behavior was conserved, regardless of differences in tree leaf area.     

Effects of thinning on soil and tree water relations, transpiration and growth in an oak forest (Quercus petraea (Matt.) Liebl.)

To quantify the effects of crown thinning on the water balance and growth of the stand and to analyze the ecophysiological modifications induced by canopy opening on individual tree water relations, we conducted a thinning experiment in a 43-year-old Quercus petraea stand by removing trees from the upper canopy level. Soil water content, rainfall interception, sap flow, leaf water potential and stomatal conductance were monitored for two seasons following thinning. Seasonal time courses of leaf area index (LAI) and girth increment were also measured. Predawn leaf water potential was significantly higher in trees in the thinned stand than in the closed stand, as a consequence of higher relative extractable water in the soil. The improvement in water availability in the thinned stand resulted from decreases in both interception and transpiration. From Year 1 to Year 2, an increase in transpiration was observed in the thinned stand without any modification in LAI, whereas changes in transpiration in the closed stand were accompanied by variations in LAI. The different behaviors of the closed and open canopies were interpreted in terms of coupling to the atmosphere. Thinning increased inter-tree variability in sap flow density, which was closely related to a leaf area competition index. Stomatal conductance varied little inside the crown and differences in stomatal conductance between the treatments appeared only during a water shortage and affected mainly the closed stand. Thinning enhanced tree growth as a result of a longer growing period due to the absence of summer drought and higher rates of growth. Suppressed and dominant trees benefited more from thinning than trees in the codominant classes.     

基于土壤水分承载力的林分密度计算与调控——以六盘山华北落叶松人工林为例

立地水分条件决定的植被承载力是干旱缺水地区森林合理经营的重要依据。考虑到干旱缺水地区的森林蒸散耗水在水分输出中占据绝对主导地位,其大小直接与叶面积指数(LAI)相关,将林冠LAI在生长季一段时间内的最大值(LAImax)作为植被承载力(LAIc)的量化指标,利用冠层分析仪(LAI-2000),在六盘山香水河小流域和叠叠沟小流域的44个华北落叶松人工林样地,实测了冠层LAI的季节动态变化,研究了生长季内LAImax与林分断面积、郁闭度、平均树高、密度等常用林分结构指标的关系。结果表明:LAImax与林分不同结构指标均呈幂函数关系,其决定系数(R2)依次为0.84、0.82、0.56、0.47,说明能同时反映林分密度和树体大小的林分断面积与林冠LAI相关最紧密。将LAImax与林分断面积的幂函数关系嵌入了林分平均胸径与林分密度和林龄关系的模型,用以描述LAImax与林龄和密度的关系,并利用样地实测数据拟合了模型参数。拟合建立的模型对所有样地的LAImax的计算值与实测值的相对误差平均为8.6%(0%20.4%),能较好地描述LAI与林龄和密度的关系。利用此模型,进一步导出了能依据给定的LAIc,简捷计算出不同林龄时的可承载林分密度的模型,从而为基于立地水分植被承载力的林分密度管理和森林多功能经营等提供技术支持。     

History of Peridermium Stem Rust of Scots Pine (Pinus sylvestris L.) in Thetford Forest, East Anglia

‘Peridermium’ stem rust (Peridermium pini (Pers.)Lev.) has been present in Thetford forest for at least 40 years,but has only become a major problem during the last decade.Survey data are presented which show that there has been a dramaticincrease in the disease in crops of Scots pine between 1964,when with less than one per cent of the trees showed symptoms,and 1979, when the figure was 10 per cent. The disease has apparentlyspread outwards from a central focus in the middle of the forest.In four plots the proportion of trees with visible symptomshas increased in five years from an average of 28 per cent to46 per cent. However, only 1–2 per cent of the trees havedied annually, and it appears that many trees with ‘deadtops’ may survive for long periods. The situation in Thetfordseems to contrast with that in north-east Scotland, where limiteddata suggest there has been no appreciable increase in diseaseincidence during recent years. There are only two records ofthe disease on Corsican pine (P. nigra var. maritima (Ait.)Melville) in Thetford.     

Temporal variation and efficiency of leaf area index in young mountain Norway spruce stand

Temporal variation of leaf area index (LAI) in two young Norway spruce stands with different densities was monitored during eight consecutive growing seasons (1998–2005). We focused on: (1) LAI dynamics and above-ground mass production of both spruce stands and their comparison, (2) leaf area duration (LADU), crop production index (CPI) and leaf area efficiency (LAE) evaluation, and (3) thinning impact on the above-mentioned parameters. Also, we tried to deduce the most effective LAI value for the Norway spruce forest investigated. The LAI values of both spruce stands showed a typical seasonal course. To describe the LAI dynamics of the stand, we recommend taking LAI measurements within short time intervals at the time of budding and needle expansion growth (i.e., in early spring) and close to the LAI peak, when the twig growth has been completed. The reason was that after reaching the seasonal maximum, no significant differences between subsequently obtained values were found in the following 2 months. Therefore, we recommend this period for the estimation of seasonally representative LAI values, enabling the comparison of various spruce stands. The maximum hemi-surface LAI value reached 12.4. Based on our results, the most effective LAI values for maximum above-ground biomass production were within the range of 10–11. We found an LAI over these values to be less effective for additional production of above-ground biomass. In forest practice, thinning intensity is mostly described by percentage of stocking reduction. We want to show that not only thinning intensity, but also the type of thinning is important information. The type of thinning significantly affected the stand above-ground biomass increment, canopy openness, stand LAI and LAI efficiency. The stimulating effect of high-type thinning was observed; the LAE as well as the CPI increased. Low-type thinning had no such effects on LAE increments compared to the high-type thinning with similar intensity.

---

     

Comparison and Dynamic Calibration between LAI Values of Larix principis-rupprechtii Plantation Determined by Canopy Scanner and Litter-fall Collection

In order to test the accuracy of the usually-used fixed calibration factor of the canopy scanner of LAI-2000 for measuring the leaf area index(LAI),a Larix principis-rupprechtii plantation was chosen in the small watershed of Xiangshuihe located at the Liupan Mountains of Ningxia Hui Autonomous Region of NW China,the LAI was measured in October 2010,a period from full canopy to the total fall of needles,by using both the LAI- 2000 and litterfall collection method.Then,a comparison was made between the LAI values determined by the litter-fall collection and that calculated based on the figures read from LAI-2000 and the fixed calibration factor(1.49).It showed that the average of LAI measurements of the 2 methods was very close,with a difference of only 5%.However,the calculated LAI from LAI-2000 was obviously higher than the true values determined by litter-fall collection when the canopy was full of needles;and obviously lower than the true value when the canopy was sparse after needle falling.The reason may be that LAI-2000 takes the projection of twigs as needles.So,a dynamic calibration factor is needed,especially in the seasons when the needle amount and the percentage of twigs projection in crown projection change quickly.Therefore,a statistic relation in a quadratic polynomial form between the 2 series of LAI data was well fitted. This relation can be used for a more accurate estimation of LAI based on the data read from the easilyused canopy scanners like LAI-2000.     

An investigation of the influence of harvesting practice on the development of blue-stain in Corsican pine logs

In September 1991, at Thetford, East Anglia and at Inverness,north-east Scotland, a comparison was initiated between blue-staindevelopment in Corsican pine logs from trees felled and processedby harvesting machine and similar logs from trees felled andtrimmed out by chain-saw. Blue-stain was assessed in discs from batches of logs destructivelysampled at 2-week intervals after felling. At the more southerlysite of Thetford, stain was first observed after 4 weeks. Themaximum recorded area was 10 per cent of the wood surface ina sample analysed after 10weeks. In contrast, stain never exceeded,1.1 per cent of the wood surface at Inverness. Amounts of stainwere significantly greater with machine-harvested logs thanwith chainsaw-harvested logs and were correlated with the amountof bark removed or loosened during harvesting. The use of spikedrollers resulted in more stain than did the use of rubber rollers. The fungi isolated from stained wood included Sphaeropsis sapinea,Potebniamyces coniferarum and Ceratocystis coerulescens. Theseare species adapted to the colonization of wood exposed throughdamage to the overlying bark     

Calibration and assessment of seasonal changes in leaf area index of a tropical dry forest in different stages of succession

A simple measure of the amount of foliage present in a forest is leaf area index (LAI; the amount of foliage per unit ground surface area), which can be determined by optical estimation (gap fraction method) with an instrument such as the Li-Cor LAI-2000 Plant Canopy Analyzer. However, optical instruments such as the LAI-2000 cannot directly differentiate between foliage and woody components of the canopy. Studies investigating LAI and its calibration (extracting foliar LAI from optical estimates) in tropical forests are rare. We calibrated optical estimates of LAI from the LAI-2000 with leaf litter data for a tropical dry forest. We also developed a robust method for determining LAI from leaf litter data in a tropical dry forest environment. We found that, depending on the successional stage of the canopy and the season, the LAI-2000 may underestimate LAI by 17% to over 40%. In the dry season, the instrument overestimated LAI by the contribution of the woody area index. Examination of the seasonal variation in LAI for three successional stages in a tropical dry forest indicated differences in timing of leaf fall according to successional stage and functional group (i.e., lianas and trees). We conclude that when calculating LAI from optical estimates, it is necessary to account for the differences between values obtained from optical and semi-direct techniques. In addition, to calculate LAI from litter collected in traps, specific leaf area must be calculated for each species rather than from a mean value for multiple species.     

华北落叶松人工林叶面积指数实测值与冠层分析仪读数值的比较和动态校正

叶面积指数(LAI)是森林生态系统的重要结构参数,用来反映植物叶量和群体生长特征[1-4],对生态系统水分和养分循环、地表和大气之间的相互作用等诸多过程都有重要影响[5-6],广泛应用于植物生态、植物生理、生态水文以及一些交叉学科的研究中,也可作为科学培育森林、评价林分质量的定量指     

Response of LAI-2000 estimates to changes in plant surface area index in a Scots pine stand

We assessed the accuracy with which the LAI-2000 plant canopy analyzer measured changes in leaf area index (LAI) and plant area index (PAI) in a 25-year-old Scots pine (Pinus sylvestris L.) stand. Stand density was 2100 stems ha(-1) and mean tree height was 8.7 m. Needle and branch areas of the stand were reduced progressively to zero by the stepwise removal of branches on all trees growing in a circular plot with a radius of 25 m. An LAI-2000 estimate was taken after each step reduction. The needle and branch surface areas removed at each step were estimated from direct measurements and were compared with the changes in the LAI-2000 estimates. Initially (before removal of branches), directly measured PAI was 5.2 (needles = 86%, branches = 8% and stems = 6%). The LAI-2000 estimate of total surface area was 66% of direct PAI and 77% of direct LAI. There was a nonlinear relationship between the LAI-2000 estimate and directly measured PAI, such that their ratio (equivalent to the clumping factor) increased from 0.66 to 1.05 with decreasing PAI. At the last measurement, when only stems were left, the LAI-2000 estimate agreed well with the direct measurement of PAI. The LAI-2000 underestimated the direct measurement of LAI at the first three steps when LAI was > 2 and the proportion of woody area was small (< 20%). However, because the LAI-2000 estimate included stem and branch areas, it overestimated the direct measurement of LAI at the last three measurements when the proportion of woody area was large (> 20%).     

Allometry and evaluation of in situ optical LAI determination in Scots pine: a case study in Belgium

We evaluated several optical methods for in situ estimation of leaf area index (LAI) in a Belgian Scots pine (Pinus sylvestris L.) stand. The results obtained were compared with LAI determined from allometric relationships established in the same stand. We found high correlations between branch cross-sectional area, diameter at breast height (DBH) and basal area as dependent variables, and leaf mass, needle area and crown projection as independent variables. We then compared LAI estimated by allometry with LAI determined by three optical methods (LAI-2000, TRAC and digital hemispherical photography) both before and after corrections for blue light scattering, clumping and non-leafy material. Estimates of stand LAI of Scots pine ranged from 1.52 for hemispherical photography to 3.57 for the allometric estimate based on DBH. There was no significant difference (alpha = 0.01) between the allometric LAI estimates and the optical LAI values corrected for blue light scattering, clumping and interception by non-leafy material. However, we observed high sensitivity of the optical LAI estimates to the various conversion factors, particularly to the clumping factor, indicating the need for caution when correcting LAI measured by optical methods.     

Leaf distribution in large trees and stands of the floodplain forest in southern Moravia

Vertical distributions of leaf dry mass (M(d)) and leaf area (A(f)) were related to relative irradiance (I(r); I(r) above the stand = 1) in closed-canopy, old-growth stands of the floodplain forest in southern Moravia composed largely of Quercus, Fraxinus and Tilia species. Foliage area and mass at any given canopy height were converted to solar equivalent leaf area (A(s)) and mass (M(s)) by multiplying actual values at a given level in the canopy by the relative irradiance at that position. Stand leaf area index (LAI) was 5 (7 including shrub and herb layer), and solar equivalent parameters reached about 25% of that amount. In all species, vertical profiles of both relative irradiance and leaf dry mass to area ratio (LMA) were sigmoidal and the two variables were linearly related. The dominant, upper canopy species had a larger proportion of solar equivalent foliage than suppressed understory species. For individual trees of all species, the upper canopy had a larger proportion of solar equivalent foliage than the lower canopy. Light compensation points at both the leaf and whole-tree level were defined according to leaf or tree position, size and structure. I conclude that optimization of A(s) for forest stands may be used as a basis for determining thinning schedules and evaluating tree survival after damage to tree crowns by various factors.     

New management options in chestnut coppices: an evaluation on ecological bases

The effects of long rotation periods and heavy low thinning on chestnut (Castanea sativa Mill.) coppices have been evaluated from a bio-ecological point of view. Aboveground tree biomass and its partitioning, bio-ecological indexes such as litter production, leaf area index (LAI), radiation regime, and growth efficiency have been analyzed for 4 years in thinned and unthinned permanent plots established in a coppice stand aged 11 years under normal rotation (NR) and in a coppice stand aged 35 years under long rotation (LR). A decrease in LAI, litter production and growth rate with the age was observed. LR showed high current annual increments (>16.0 m³ ha⁻¹ per year and 8.0 Mg ha⁻¹ per year for volume and aboveground biomass, respectively). Only slight differences in growth efficiency were observed. The adoption of heavy thinning (one third of basal area removal) affected stocking and determined significant differences in the light regime below the canopy, amount of gaps in canopy cover and LAI values, particularly in the years immediately after thinning, whereas slight differences were observed in growth efficiency. Nevertheless, chestnut showed a good aptitude, more evident in the younger stand, to re-build a homogeneous canopy cover: only a few years after thinning, canopy cover characteristics of thinned plots were similar to those of control plots and differences were not significant. Growth and increments in thinned plots were practically the same as in control plots, a consequence of consistently higher performances of released trees in the thinned plots. The results concur to a positive evaluation of a cultivation system based on long rotation periods and heavy thinnings, not only for the obtainable revenue, but especially from a bio-ecological point of view and make it a valid alternative either to abandonment or traditional over-exploitation of chestnut coppice stands.     

Wind Measurements in a Pine Forest During a Destructive Gale

Anemometer readings were recorded during a gale which blew downsome of the trees in an even-aged 16 m stand of Pinus sylvestrison sandy soil at Thetford forest. Gusts at the top of the canopyattained 17•5 m/sec. Wind profiles agreed well with thetheoretical logarithmic profile above the canopy and the exponentialprofile below. During the gale the zero plane displacement androughness length values were similar to those at lower windspeeds. It is therefore possible to estimate wind profiles andthe forces to which a tree in the crop is subjected from measurementsof wind speed at a single point within or above the forest.The wind speeds which blew trees down were much lower than thosepredicted as necessary from ‘tree pulling’ studiesin the same forest. This discrepancy suggests that any considerationof windthrow must take into account the effects of tree vibrationon the deterioration in the strength of the root-soil complex.     

保定市常见城市绿化树种对PM2.5滞纳能力的研究

为了探究绿化树种对改善城市污染的影响,选取保定市11种常见城市绿化树种,对其滞纳PM2.5能力进行了探讨。利用冠层分析仪(LAI-2000)、叶面积扫描仪和气溶胶再发生器(QRJZFSQ—I)测定各树种的叶面积指数、单位叶面积PM2.5滞纳量,并计算单位绿地面积PM2.5滞纳量,结果表明:叶面积指数、单位叶面积PM2.5滞纳量、单位绿地面积PM2.5滞纳量3个指标,都是针叶树大于阔叶树,说明针叶树对PM2.5的滞纳能力高于阔叶树;针叶树、阔叶树叶面积指数范围分别为2.172 1~3.026 3、0.873 8~2.120 5,单位叶面积PM2.5滞纳量范围分别为0.53~1.39mg/m~2、0.11~0.34mg/m~2,单位绿地面积PM2.5滞纳量范围分别为1.151 2~4.206 6mg/m~2、0.108 8~0.622 8mg/m~2。     

Why does leaf nitrogen decline within tree canopies less rapidly than light? An explanation from optimization subject to a lower bound on leaf mass per area

A long-established theoretical result states that, for a given total canopy nitrogen (N) content, canopy photosynthesis is maximized when the within-canopy gradient in leaf N per unit area (N(a)) is equal to the light gradient. However, it is widely observed that N(a) declines less rapidly than light in real plant canopies. Here we show that this general observation can be explained by optimal leaf acclimation to light subject to a lower-bound constraint on the leaf mass per area (m(a)). Using a simple model of the carbon-nitrogen (C-N) balance of trees with a steady-state canopy, we implement this constraint within the framework of the MAXX optimization hypothesis that maximizes net canopy C export. Virtually all canopy traits predicted by MAXX (leaf N gradient, leaf N concentration, leaf photosynthetic capacity, canopy N content, leaf-area index) are in close agreement with the values observed in a mature stand of Norway spruce trees (Picea abies L. Karst.). An alternative upper-bound constraint on leaf photosynthetic capacity (A(sat)) does not reproduce the canopy traits of this stand. MAXX subject to a lower bound on m(a) is also qualitatively consistent with co-variations in leaf N gradient, m(a) and A(sat) observed across a range of temperate and tropical tree species. Our study highlights the key role of constraints in optimization models of plant function.     

Estimating leaf area index of mature temperate forests using regressions on site and vegetation data

Canopy gap fraction and leaf area index (LAI) were measured using hemispherical photography in 91 mature forests across Switzerland, including coniferous, broadleaved and mixed stands. The gap fraction and LAI derived from five photographs per site could be reproduced with a high coefficient of determination (R² > 0.7) by regression against simple stand parameters obtained from vegetation surveys: coverages of the tree, shrub and herb layers, and tree height. The method appeared to be robust across the different types of forests. Applied to 981 sites across Switzerland, the regression model produced LAI values ranging from 1.4 to 6.7. These predictions were compared with site variables not included in the regression. LAI appeared limited by the altitude, with maximal values decreasing by one third from 400 to 2000 m above see level. Water availability was also clearly a limitation at sites with a negative water balance, i.e. where the yearly potential evapotranspiration exceeded the precipitation. High or low values of a humidity index based on the ground vegetation also corresponded to a limitation of the LAI, with shorter trees at dry sites and more open canopies at wet sites. Compared to optical measurements (including hemispherical photography), our regression method is fast and inexpensive. Such an approach appears very promising for obtaining reliable estimates of LAI for many sites with low costs. These estimates can then be fed into process models at the stand level.     

A comparison of red spruce and balsam fir shoot structures

I compared the shoot structures of high-elevation red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea (L.) Mill.). Needle widths, thicknesses and perimeters were measured to estimate total leaf areas from measured projected leaf areas. Measured needle perimeter/needle width ratios differed significantly from estimated ratios that assumed needles were either rhomboidal or elliptical in cross section. The vertical and horizontal silhouette shoot area to total leaf area ratios (STAR(v) and STAR(h)) of the two species were negatively correlated with needle packing and canopy height. Red spruce had higher values of STAR(v) than balsam fir at each canopy height, but STAR(v) declined with canopy height at a similar rate in the two species. The STAR(h) values of the two species did not differ significantly at a given canopy height. Needle packing increased with canopy height at the same rate in the two species. Needle weight increased in red spruce and decreased in balsam fir with increased needle packing, but showed no significant dependence on canopy height. Red spruce had higher values of STAR(h) than balsam fir at low values of needle packing, but STAR(h) values converged at high values of needle packing. The generally comparable values of STAR, along with similar needle diameters, may imply that red spruce and balsam fir have similar collection efficiencies of wet and dry particles. Measurements of STAR may be used to estimate leaf area indices (LAI) more accurately when using indirect techniques.     

Response of Douglas-fir leaf area index and litterfall dynamics to Swiss needle cast in north coastal Oregon, USA

Sources of variation in leaf area index (LAI; m² of projected leaf area per m² of ground area) and its seasonal dynamics are not well known in managed Douglas-fir stands, despite the importance of leaf area in forecasting forest growth, particularly in stands impacted by insects or disease. The influence of Swiss needle cast (SNC) on coastal Douglas-fir (Pseudotsuga menziesii var. menziesii [Mirb] Franco) LAI and litterfall dynamics was quantified by destructively sampling 122 stems from 36 different permanent plots throughout north coastal Oregon, USA, and by monitoring litterfall for 3 years in 15 of these plots. LAI, total annual litterfall, and the seasonal distribution of foliage and fine woody litterfall were all influenced by stand structural attributes, physiographic features, and SNC severity. Mean LAI in this study was 5.44 ± 2.16. The relatively low LAIs were attributed primarily to the effects of SNC on foliage retention, and secondarily to its direct measurement by hierarchical foliage sampling in contrast to indirect measurement by light interception or tree allometry. For a given stand structure and SNC severity, LAI was 36% greater in the fall after current year foliage was fully developed and older aged classes had not yet senesced. Annual litterfall expressed as a proportion of LAI at the start of the growing season varied from 0.13 to 0.53 and declined with increasing initial LAI. SNC also shifted more of the annual foliage litterfall to earlier in the spring. Fine woody litterfall experienced a different seasonal shift as the peak occurred later in the year on sites with high SNC, but this only occurred on northerly aspects. Defoliation from the endemic SNC pathogen can drastically reduce LAI and change both total and seasonal foliage litterfall patterns.     

Age-related effects on leaf area/sapwood area relationships,canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge,Germany

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.