Estimating foliage biomass in Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) plots

Dynamic decomposition models are needed to estimate changes in the carbon stock of boreal soil because these changes are difficult to measure directly. An important aboveground carbon flux to the soil is foliage litterfall. To estimate this flux, both the amount and the turnover rate of the foliage biomass component must be known. Several methods for estimating foliage biomass of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.), including biomass equations and biomass expansion factors (BEFs), were compared with predicted foliage biomass based on forest inventory plot-level measurements. Measured foliage biomass was up-scaled from the branch-level to the plot-level by combining forest inventory variables (diameter, height, height at the crown base and crown base diameter) based on the assumptions of pipe model theory. Combining the foliage biomass: cross-sectional area ratio with the forest inventory variables provided accurate estimates of foliage biomass at the plot-level for plots in southern Finland. The results emphasize the need to test biomass equations with independent data, especially when the equations applied are based on neighboring regions.     

Quantifying the effect of pine mistletoe on the growth of Scots pine

Mistletoe infection results in substantial growth losses in mistletoe‐infected forests. This study reports and evaluates the results of retrospective analyses of radial growth of Scots pine (Pinus sylvestris) in relation to the level of infection of pine mistletoe (Viscum album ssp. austriacum). A total of 43 Scots pine trees were destructively sampled from different sites. Of these trees, 14 were uninfected and 29 were infected. Infection classes were determined using six‐class dwarf mistletoe rating system (DMRS). All needle and mistletoe biomass were removed completely and weighed for each sampled tree. Subsamples from needles and all mistletoe biomass were taken to the laboratory for oven‐dried weight determinations. Five‐cm‐thick wood discs were cut from the stem at the breast height (1.3 m) to determine annual basal area increment for the last 25 years. In addition to DMRS, new infection classes were created using mistletoe‐to‐needle biomass (MB/NB) ratio. The results showed that the radial growth losses could be as much as 41% to 64% at different infection levels. The rate of growth loss in relation to DMRS and MB/NB ratio was similar, but with a larger variability in DMRS values. The results showed that both DMRS rating and MB/NB ratio seem to be important for quantifying growth loss on Scots pine trees infected with mistletoe. The results of this study can also be invaluable in modelling the effects of mistletoe on the growth of Scots pine trees.     

Needle and stem wood production in Scots pine (Pinus sylvestris) trees of different age,size and competitive status

We studied effects of tree age, size and competitive status on foliage and stem production of 43 Scots pine (Pinus sylvestris L.) trees in southern Finland. The tree attributes related to competition included foliage density, crown ratio and height/diameter ratio. Needle mass was considered to be the primary cause of growth through photosynthesis. Both stem growth and foliage growth were strongly correlated with foliage mass. Consequently, differences in growth allocation between needles and stem wood in trees of different age, size, or position were small. However, increasing relative height increased the sum of stem growth and foliage growth per unit foliage mass, indicating an effect of available light. Suppressed trees seemed to allocate more growth to stem wood than dominant trees, and their stem growth per unit foliage mass was larger. Similarly, trees in dense stands allocated more growth to stem wood than trees in sparse stands. The results conformed to the pipe model theory but seemed to contradict the priority principle of allocation.     

Impacts of thinning on structure,growth and risk of crown fire in a Pinus sylvestris L. plantation in northern Spain

We studied the combined effects of thinning on stand structure, growth, and fire risk for a Scots pine thinning trial in northern Spain 4 years following treatment. The thinning treatments were: no thinning, heavy thinning (32–46% of basal area removed) and very heavy thinning (51–57% of basal area removed). Thinning was achieved via a combination of systematic and selective methods by removing every seventh row of trees and then by cutting suppressed and subdominant trees in the remaining rows (i.e., thinning from below). Four years after thinning, mean values and probability density distributions of stand structural indices showed that the heavier the thinning, the stronger the tendency towards random tree spatial positions. Height and diameter differentiation were initially low for these plantations and decreased after the 4-year period in both control and thinned plots. Mark variograms indicated low spatial autocorrelation in tree diameters at short distances. Diameter increment was significantly correlated with the inter-tree competition indices, and also with the mean directional stand structural index. Two mixed models were proposed for estimating diameter increment using a spatial index based on basal area of larger trees (BALMOD) in one model versus spatial competition index by Bella in the other model. As well, a model to estimate canopy bulk density (CBD) was developed, as this variable is important for fire risk assessment. Both heavy and very heavy thinning resulted in a decrease of crown fire risk over no thinning, because of the reduction in CBD. However, thinning had no effect on the height to crown base and thus on the flame length for torching. Overall, although thinning did not increase size differentiation between trees in the short term, the increase in diameter increment following thinning and the reduction of crown fire risks support the use of thinning. Also, thinning is a necessary first step towards converting Scots pine plantations to more natural mixed broadleaved woodlands. In particular, the very heavy thinning treatment could be considered a first step towards conversion of overstocked stands.     

The effects of pine mistletoe (Viscum album subsp. austriacum) on the growth of Scots pine and Crimean pine in Turkey

In this study, the effect of pine mistletoe (Viscum album subsp. austriacum) on basal area increment of Crimean pine and Scots pine was investigated. Dendrochronological data were collected from 223 (71 uninfected and 152 infected) Crimean pines and 195 (77 uninfected and 118 infected) Scots pines located in Kastamonu province of Turkey in 2014. Infected sample trees were classified as light, moderate or severe infection levels. Growth trends and basal area increment loses were compared between uninfected and infected trees for the periods of the last 10, 20 and 30 years. In addition, infection status of forest stands was investigated using temporary sample plots; 27 plots in Crimean pine stands and 26 plots in Scots pine. Results demonstrated that basal area increments were negatively affected by pine mistletoe for both species. Mean basal area increment losses of infected trees for the last decade were determined as 24% for Scots pine and 26% for Crimean pine. Basal area increment losses varied by infection levels (light, moderate and severe) as follows: 25%, 20% and 28% for Scots pines and 20%, 32% and 9% for Crimean pines. Scots pine stands were more severely infected by pine mistletoe than Crimean pine stands. There were negative correlations between number of infected trees and stand density for both species, while positive correlation was detected between the number of infected trees and mean diameter for Scots pine. The results of this study indicate that the pine mistletoe infection has negative effect on radial growth of Scots pine and Crimean pine trees. The results can be an important contribution to the forest management and protection activities in mistletoe-infected stands.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Sapwood Basal Area and Needle Mass of Scots Pine (Pinus sylvestris L.) Trees in Central Sweden

The relationship between foliage mass or area and sapwood basalarea is of potential interest in the quest for cheap and simplemethods of estimating foliage biomass. The sapwood basal area and the needle dry weight were obtainedby destructive sampling for 153 Scots pine trees (Pinus sylvestrisL.) from 16 stands in central Sweden. The regression relationshipbetween needle mass and sapwood basal area was strong for singlestands, but it did not seem appropriate to aggregate the materialinto one overall regression, having sapwood basal area as theonly independent variable. It was shown that the needle biomassper unit of sapwood basal area varied with mean annual ringwidth in the sapwood. This information was used in an improvedoverall model.     

Aboveground biomass of three conifers in the Qianyanzhou plantation, Jiangxi Province, China

Regressive models of the aboveground biomass for three conifers in subtropical China—slash pine (Pinus elliottii), Masson pine (P. massoniana) and Chinese fir (Cunninghamia lanceolata)—were established. Regression analysis of leaf biomass and total biomass of each branch against branch diameter (d), branch length (L), d ³ and d ² L was conducted with functions of linear, power and exponent. A power law equation with a single parameter (d) was proved to be better than the rest for Masson pine and Chinese fir, and a linear equation with parameter (d ³) is better for slash pine. The canopy biomass was derived by adopting the regression equations to all branches of each individual tree. These kinds of equations were also used to fit the relationship between total tree biomass, branch biomass, foliage biomass and tree diameter at breast height (D), tree height (H), D ³ and D ² H, respectively. D ² H was found to be the best parameter for estimating total biomass. However, for foliage biomass and branch biomass, both parameters and equation forms showed some differences among species. Correlations were highly significant (P<0.001) for foliage biomass, branch biomass and total biomass, among which the equation of the total biomass was the highest. With these equations, the aboveground biomass of Masson pine forest, slash pine forest and Chinese fir forest were estimated, in addition to the allocation of aboveground biomass. The above-ground biomass of Masson pine forest, slash pine forest and Chinese fir forest was 83.6, 72.1 and 59 t/hm² respectively, and the stem biomass was more than the foliage biomass and the branch biomass. The underground biomass of these three forests which estimated with others’ research were 10.44, 9.42 and 11.48 t/hm², and the amount of carbon-fixed were 47.94, 45.14 and 37.52 t/hm², respectively. __________ Translated from Chinese Journal of Applied Ecology, 2006, 17(8): 1382–1388 [译自: 应用生态学报]     

气候敏感的马尾松生物量相容性方程系统研建

【目的】构建气候敏感的马尾松生物量相容性方程系统,分析气候因子对马尾松各分项生物量的影响,为森林碳汇监测和森林可持续经营提供技术支撑。【方法】基于150株马尾松单木生物量数据,采用非线性联立方程组法构建气候敏感的马尾松生物量相容性方程系统,各分项生物量(干材、干皮、树枝、树叶和地上总生物量)选用以直径和树高为自变量的二元生物量方程作为基础模型,利用一阶交叉验证法对所构建的生物量相容性方程系统进行评价。【结果】与传统未考虑气候因子的各分项生物量模型相比,气候敏感的马尾松生物量相容性方程系统预测精度明显提高,且该生物量相容性方程系统可定量描述不同气候带亚区生物量的差异程度,保证干材、干皮、树枝和树叶与地上总生物量相容。【结论】气候敏感的马尾松生物量相容性方程系统能有效分析气候因子对各分项生物量的影响,可应用于其他树种的生物量预估。     

Derivation of stem taper from the pipe theory in a carbon balance framework

A dynamic tree growth model is described. The model derives the development of stem taper and vertical distribution of branch basal area from the pipe model, assuming that reuse of active pipes is regulated by foliage dynamics in a vertically explicit crown with a foliage distribution of constant shape. Based on empirical findings, the pipe model was modified slightly to allow the foliage/sapwood ratio to vary as a function of distance from the treetop. Growth was derived from carbon balance in a stand of different size trees that may shade each other. The model was applied to old and middle-aged trees growing in dense and sparse stands of Scots pine for which stand-level measurements are available as a chronosequence, but individual trees have been measured only once. Measured trees were compared with corresponding simulated trees for stem taper and vertical distribution of branch basal area. The results indicated that the pipe model assumptions, combined with a model of tree growth, are capable of producing realistic predictions of the vertical distribution of stem and branch diameter in trees of different sizes in the stand. A comparison of the results with a simple form of the uniform stress theory showed good agreement between the two models. However, a significant difference was found between the measured relative contribution of heartwood to total stem diameter and the predicted share of disused pipes in the stem. A possible explanation for this discrepancy is that the transition from sapwood to heartwood is gradual rather than abrupt as assumed in the model. A modification of the pipe model to incorporate a gradual transition is outlined.     

Aboveground biomass equations for individual trees of <Emphasis Type="Italic">Cryptomeria japonica</Emphasis>, <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> and <Emphasis Type="Italic">Larix kaempferi</Emphasis> in Japan

Generic equations are proposed for stem, branch and foliage biomass of individual trees in even-aged pure stands of Cryptomeria japonica, Chamaecyparis obtusa and Larix kaempferi. Biomass data was collected from a total of 1,016 individual trees from 247 stands throughout Japan, and five regression models were assessed by root mean square error, mean bias, fit index (FI), and AIC. The results show that a power equation using diameter at breast height (dbh) and height is the most suitable for all species and components. This equation is more accurate than the familiar power equation that uses ‘dbh² height’, and it expresses the greater volume of branch and foliage mass of trees with a lower height/diameter ratio. A power equation using dbh is more reasonable for models with dbh as the only independent variable and more accurate than a power equation using ‘dbh² height’ for estimating branch and foliage mass. Estimating error for branch and foliage mass is larger than that for stem mass, but the entire aboveground biomass can be estimated with an error of less than 19%, except in the case of small trees with dbh less than 10 cm.     

Canopy fuel characteristics in relation to crown fire potential in pine stands: analysis, modelling and classification

Crown fire occurrence and subsequent crown fire behaviour are strongly dependent on canopy fuel characteristics, especially canopy fuel load (CFL), canopy bulk density (CBD) and canopy base height (CBH). Therefore, quantification of such variables is required for the appropriate selection of silvicultural treatments aimed at reducing susceptibility to crown fire. Data from the IV Spanish National Forest Inventory and individual tree biomass dry weight equations were used to estimate the canopy fuel characteristics of four representative types of pine stands in north-western Spain. Probability of crown fire initiation and crown fire rate of spread were simulated by using the mean surface fuel load observed for each type of pine in this area and assuming different burning conditions. The results indicate that a 22.13 % of the sample plots analysed showed a rather high potential for active crown fire spread under moderate burning conditions, and this value increases to 69.27 % under extreme burning conditions. Equations relating the canopy fuel characteristics to common stand variables (stand density, basal area and dominant height) were fitted simultaneously for each pine, and weighting factors for heteroscedasticity were included. The models explained more than 93.90, 74.70 and 69.42 % of the observed variability in CFL, CBD and CBH, respectively. Basal area was the most important variable for estimating CFL and CBD while dominant height explained most of the observed variability in CBH. The use of the fitted equations together with existing dynamic growth models and fire management decision support systems will enable assessment of the crown fire potential associated with different silvicultural alternatives used in these types of pine stands.     

Integrated individual tree biomass simultaneous equations for two larch species in northeastern and northern China

Forest biomass estimation at large scale has become an important topic in the background of facing global climate change, and it is fundamental to develop individual tree biomass equations suitable for large-scale estimation. Based on the measured data of biomass components and stem volume from 100 sample trees of two larch species (Larix gmelinii and L. principis-rupprechtii) in northeastern and northern China, an integrated equation system including individual tree biomass equations, stem volume equation and height–diameter regression model were constructed using the dummy variable model and error-in-variable simultaneous equations. In the system, all the parameters of equations were estimated simultaneously, so that the aboveground biomass equation was compatible to stem volume equation and biomass conversion factor (B_CF) function; the belowground biomass equation was compatible to root-to-shoot ratio (R_SR) function; and stem wood, stem bark, branch and foliage biomass equations were additive to aboveground biomass equation. In addition, the system also ensured the compatibility between one- and two-variable models. The results showed that: (1) whether aboveground biomass equations or belowground biomass equations and stem volume equations, the estimates for larch in northeastern China were greater than those in northern China; (2) B_CF of a larch tree decreased with the growing diameter while R_SR increased with the growing diameter; (3) the proportion of stem wood biomass to aboveground biomass increased with the growing diameter while those of stem bark, branch, and foliage biomass decreased.     

Structural development of pinus sylvestris stands with varying initial density: A simulation model

The growth and structural development of Scots pine (Pinus sylvestris L.) trees growing at different spacing was simulated using a model based on the dry matter production per needle biomass unit and its allocation to needles, branches and stem. Special emphasis was given to the effect of stand density on the growth of the crown system and its implications on branchiness and timber quality. The simulations showed that the needle biomass culminates considerably earlier than the branch biomass with a time lag inversely related to the stand density. The lengths of living and dead crown were also inversely related to stand density. The resulting differences in branchiness were especially obvious in the early development of the tree stands. In the long run these differences tend to disappear, indicating equal external branchiness independently of the initial spacing for mature stands of Scots pine. The internal branchiness, however, was particularly sensitive to the initial spacing.     

Thinning alters crown dynamics and biomass increment within aboveground tissues in young stands of Chamaecyparis obtusa

The stand density of a forest affects the vertical distribution of foliage. Understanding the dynamics of this response is important for the study of crown structure and function, carbon-budget estimation, and forest management. We investigated the effect of tree density on the vertical distribution of foliage, branch, and stem growth, and ratio of biomass increment in aboveground tissues; by monitoring all first-order branches of five trees each from thinned and unthinned control stands of 10-year-old Chamaecyparis obtusa for four consecutive years. In the control stand, the foliage crown shifted upward with height growth but the foliage quantity of the whole crown did not increase. In addition, the vertical distribution of leaf mass shifted from lower-crown skewed to upper-crown skewed. In the thinned stand in contrast, the foliage quantity of individual crowns increased two-fold within 4 years, while the vertical distribution of leaf mass remained lower-crown skewed. The two stands had similar production rates, numbers of first-order branches per unit of tree height, and total lengths of first-order branches. However, the mortality rate of first-order branches and self-pruning within a first-order branch were significantly higher in the control stand than in the thinned stand, which resulted in a higher ratio of biomass increment in branch. Thinning induced a higher ratio of biomass increment in foliage and lower in branch. The increased foliage quantity and variation in ratio of biomass increment after thinning stimulated stem growth of residual trees. These results provide information that will be useful when considering thinning regimes and stand management.     

Effect of pruning on nitrogen dynamics within crowns of Pinus radiata

Stem injection of (15)N-labeled ammonium sulfate was used to determine effects of pruning on canopy nitrogen dynamics in open-grown Pinus radiata D. Don in New Zealand. Trees were planted in July 1990 and the isotope introduced in December 1994. Tree crowns were divided into three zones: base section, from which branches of pruned trees were removed; mid section, between the pruned zone and the height of the trees at the start of the year in which they were pruned; and top section, which grew predominantly after the isotope was applied. Pruning removed 32% of the green crown length, representing 75% of foliage biomass. Needles were sampled from each region of the crown until July 1996. Branch growth was used to predict foliage biomass for each sampling occasion. Approximately 45% of the applied isotope was recovered from needles sampled in December 1994 (1 week after application and immediately before pruning), two-thirds of which occurred in needles in the base section. Thereafter, changes in isotope content of needles in the base section of unpruned trees largely reflected foliage biomass fluctuations and dilution of the isotope by continued uptake from the unlabeled soil nitrogen pool. Recovery of isotope in needles from the mid-crown section increased by 58 and 86% from December 1994 to July 1995 in control and pruned trees, respectively. Within this crown section, there was evidence of isotope translocation from old to new needles, with both isotope dilution and efflux observed in the needle cohorts that had been present at the time the isotope was applied. Therefore, isotope dynamics did not reflect the dynamics of the total nitrogen pool in the mid-crown section. By July 1996, a small proportion of the applied isotope was recovered from the new foliage formed in the top section of the crown. Within the top section, isotope dynamics closely matched total nitrogen fluxes. Pruning the lower crown did not affect nitrogen dynamics elsewhere in the crown for the following 18 months.     

A basal area increment model for individual trees growing in even- and uneven-aged forest stands in Austria

Because of the gradual shift from pure even-aged forest management in central Europe, existing yield tables are becoming increasingly unreliable for forest management decisions. Individual tree-based stand growth modeling can make accurate stand growth predictions for the full range of conditions between pure even-aged and mixed-species uneven-aged stands. The central model in such a simulator is basal area increment for individual trees. Spatial information is not needed, and age and site index are intentionally not used to gain generality for all possible stand conditions. A basal area increment model is developed for all the main forest species in Austria: spruce (Picea abies), fir (Abies alba), larch (Larix decidua), Scots pine (Pinus sylvestris), black pine (Pinus nigra), stone pine (Pinus cembra), beech (Fagus silvatica), oak (Quercus robur, Quercus petraea and Quercus cerris), and for all other broadleaf species combined. The Austrian National Forest Inventory provided 5-year basal area increment from 44 761 remeasured trees growing on 5416 forested plots in the 1980s. This large sample is representative of forest conditions and forest management practices throughout Austria and therefore provides an excellent data base for the development of an increment model. The resulting increment model explained from 20 to 63% of the variation for all nine species and from 33 to 63% of the variation if the minor species Pinus cembra is excluded. These results compared quite closely with those of Wykoff for mixed conifer stands in the Northern Rocky Mountains. In the Austrian model, size variables (breast height diameter and crown length) accounted for 14–47% of the variation in basal area increment, depending on tree species. The best competition measure was the basal area of larger trees, which provides a tree-specific measure of competition without requiring spatial information; crown competition factor provided only minor improvement. Competition variables accounted for 9% of the variation on average, and up to 15% for some species. Topographic factors (elevation, slope, aspect) explained up to 3% of the variation, as did soil factors. Remaining site factors; such as vegetation type and growth district accounted for a maximum of 3% of the variation in increment. In total, site factors explained from 2 to 6% of the variation. Even though site factors account for a small percentage of the variation, they are not only significant, but serve to localize a particular prediction. These species-specific interrelationships between basal area increment and the various size, competition, and site varibles correspond quite well with ecological expectations and silvicultural understanding of these species in Austria. Because the sample base is so strong, the resulting growth models can be recommended not only for all of Austria but for surrounding regions with similar growth conditions.     

The effect of different levels of green pruning on the diameter growth of Pinus sylvestris L.

It was hypothesized that maximum living crown removal can be determined from experimental data using linear statistical models. Pruning experiments were carried out in four Scots pine (Pinus sylvestris L.) stands. Zero to nine living branch whorls were removed in three stands in southern Finland, and 0–4.5 m of the living crown was removed in a stand in northern Finland. Only dead branch whorls were removed from the control trees. A total of 517 trees were analyzed. Diameter increment was measured five years after pruning. The growth reduction was statistically significant when 40% or more of the living crown was removed by pruning. According to the linear models, green pruning decreased diameter growth by 0 to 33% at defoliation levels below 40%. This loss in diameter growth is, from an economical point of view, compensated by the knotfree timber resulting from pruning 2–3 years earlier.     

Comparison of biomass equations for planted vs. natural loblolly pine stands of sawtimber size

Equations predicting biomass of components of sawtimber-size trees in a near-maturity loblolly pine (Pinus taeda L.) plantation were compared to similar equations for an uneven-aged natural loblolly pine stand. Combined analysis of the two sites revealed that curves estimating total tree, stem wood, stem bark, branches, and foliage + branches weights were significantly different, while curves predicting biomass for total stem and foliage were similar. Biomass equations differ because of variations in tree component ratios and taper associated with site and stand conditions.     

Carbon budget for Scots pine trees: effects of size, competition and site fertility on growth allocation and production

Time series of carbon fluxes in individual Scots pine (Pinus sylvestris L.) trees were constructed based on biomass measurements and information about component-specific turnover and respiration rates. Foliage, branch, stem sapwood, heartwood and bark components of aboveground biomass were measured in 117 trees sampled from 17 stands varying in age, density and site fertility. A subsample of 32 trees was measured for belowground biomass excluding fine roots. Biomass of fine roots was estimated from the results of an earlier study. Statistical models were constructed to predict dry mass (DW) of components from tree height and basal area, and time derivatives of these models were used to estimate biomass increments from height growth and basal area growth. Biomass growth (G) was estimated by adding estimated biomass turnover rates to increments, and gross photosynthetic production (P) was estimated by adding estimated component respiration rates to growth. The method, which predicts the time course of G, P and biomass increment in individual trees as functions of height growth and basal area growth, was applied to eight example trees representing different dominance positions and site fertilities. Estimated G and P of the example trees varied with competition, site fertility and tree height, reaching maximum values of 22 and 43 kg(DW) year(-1), respectively. The site types did not show marked differences in productivity of trees of the same height, although height growth was greater on the fertile site. The G:P ratio decreased with tree height from 65 to 45%. Growth allocation to needles and branches increased with increasing dominance, whereas growth allocation to the stem decreased. Growth allocation to branches decreased and growth allocation to coarse roots increased with increasing tree size. Trees at the poor site allocated 49% more to fine roots than trees at the fertile site. The belowground parts accounted for 25 to 55% of annual G, increasing with tree size and decreasing with site fertility. Annual G and P per unit needle mass varied over the ranges 1.9-2.4 and 3.5-4.0 kg(DW) kg(-1), respectively. The relationship between P and needle mass in the example trees was linear and relatively independent of competition, site fertility and age.