Effects of varying crop load on photosynthesis, dry matter production and partitioning of Crispin/M.27 apple trees

Fruit load was altered by flower thinning on three- and four-year-old, field-grown apple trees. Increasing fruit load led to increases in dry matter production per unit leaf area and partitioning to fruit and to decreases in fruit size, percentage fruit dry matter, dry matter partitioning to new shoot growth, thickening of existing woody tissue and root growth. Flower bud production for the following spring was also negatively affected by an increase in fruit load. Leaf photosynthesis was increased in cropping trees in July and August at the time of maximum fruit dry weight increase. Calculated light interception was linearly related to leaf area. The efficiency of conversion of intercepted photosynthetic active radiation to dry matter energy equivalents was 3.3% in heavily cropping trees and 1.8% in non-cropping trees. Total dry matter production was linearly related to both leaf area and light interception, but the variance accounted for by the regression was more than doubled if fruit dry matter or fruit number was included in the regression.     

Light modification in a developing silvopastoral system in the UK: a quantitative analysis

Transmitted photosynthetically active radiation (PAR) was measured in eight-year-old silvopastoral stands at three different locations in the UK by using PAR sensors randomly distributed between four adjacent trees planted in a square pattern. Two densities (100 stems.ha^-1 and 400 stems.ha^-1 were compared. The annual carrying capacity (live weight of the animals that a plot supports) was also recorded as an indirect figure of the overall agricultural production of the system. The silvopastoral plots received less radiation than the agricultural plot without trees, the 400 stems.ha^-1 treatment being more shaded than the 100 stems.ha^-1. Over the measurement period, on some special days, a maximum of 30% of the incoming radiation was intercepted on the 400 stem.ha^-1 treatment with the largest trees. The amount of light intercepted was highly correlated with the type of weather and the leaf area development of the trees: the brighter the day and more expanded the leaf area, the higher was the proportion of radiation intercepted by the tree. However, over the measuring period, a small amount of radiation was intercepted by the tree canopies. As a consequence, no differences were observed in the annual carrying capacity. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of crown development on leaf irradiance,leaf morphology and photosynthetic capacity in a peach tree

The three-dimensional (3-D) architecture of a peach tree (Prunus persica L. Batsch) growing in an orchard near Avignon, France, was digitized in April 1999 and again four weeks later in May 1999 to quantify increases in leaf area and crown volume as shoots developed. A 3-D model of radiation transfer was used to determine effects of changes in leaf area density and canopy volume on the spatial distribution of absorbed quantum irradiance (PAR(a)). Effects of changes in PAR(a) on leaf morphological and physiological properties were determined. Leaf mass per unit area (M(a)) and leaf nitrogen concentration per unit leaf area (N(a)) were both nonlinearly related to PAR(a), and there was a weak linear relationship between leaf nitrogen concentration per unit leaf mass (N(m)) and PAR(a). Photosynthetic capacity, defined as maximal rates of ribulose-1,5-bisphosphate carboxylase (Rubisco) carboxylation (V(cmax)) and electron transport (J(max)), was measured on leaf samples representing sunlit and shaded micro-environments at the same time that the tree crown was digitized. Both V(cmax) and J(max) were linearly related to N(a) during May, but not in April when the range of N(a) was low. Photosynthetic capacity per unit N(a) appeared to decline between April and May. Variability in leaf nitrogen partitioning between Rubisco carboxylation and electron transport was small, and the partitioning coefficients were unrelated to N(a). Spatial variability in photosynthetic capacity resulted from acclimation to varying PAR(a) as the crown developed, and acclimation was driven principally by changes in M(a) rather than the amount or partitioning of leaf nitrogen.     

Crown area allometries for estimation of aboveground tree biomass in agricultural landscapes of western Kenya

Classical allometries determine biomass from measurements of diameter at breast height or volume. Neither of these measurements is currently possible to be derived directly from remote sensing. As biomass estimates at larger scales require remotely sensed data, new allometric relations are required using crown area and/or tree height as predictor of biomass, which can both be derived from remote sensing. Allometric equations were developed from 72 trees semi-randomly selected for destructive sampling in three 100?km² sentry sites in western Kenya. The equations developed fit the data well with about 85?% of the observed variation in aboveground biomass explained by crown area. Addition of height and wood density as second predictor variables improved model fit by 6 and 2?% and lowered the relative error by 7 and 2?%, respectively. The equation with crown area in combination with height and wood density estimated representative aboveground biomass carbon to be about 20.8?±?0.02?t?C?ha^?1; which is about 19?% more than the amount estimated using an allometry with diameter at breast height as predictor. These results form the basis for a new generation of allometries using crown area as a predictor of aboveground biomass in agricultural landscapes. Biomass predictions using crown area should be supported by height and wood density and the application of crown area equations for remote sensing based up-scaling should consider crown interactions with competing or coexisting neighboring trees.     

杨树人工林冠层光合辐射分布的研究

通过对集约经营与粗放经营杨树人工林冠层上部,下部太阳总辐射和光合有效辐射（ＰＡＲ）的实测数据进行分析,建立了其相互转换的关系方程;研究了ＰＡＲ的日、季变化特征;根据Ｂｅｅｒ－Ｌａｍｂｅｒｔ方程,结合林分生长季内的叶面积动态及林冠各层次的叶面积指数（ＬＡＩ）,计算出生长季内各天的消光系数（Ｋ）及任一时刻冠层内不同深度的光分布,并对ＰＡＲ的透过率与林分消光系数Ｋ及累积叶面积指数之间的相互关系进行了研究     

Effects of partial defoliation on closed canopy Eucalyptus globulus Labilladière: Growth, biomass allocation and carbohydrates

Herbivory caused by leaf-eating insects continues to be a severe risk to forest trees and forest stands. Besides quantifying the extent of defoliation, the quantification of the trees’ response to the loss of biomass is a challenge to plant ecologists and foresters alike, and an important precondition for the application of appropriate silvicultural measures. While many defoliation studies target small trees as model systems, little is known about the effect of defoliation on larger trees. In the present study, we investigated the effects of 45% removal of leaf area on growth, biomass allocation and carbohydrates of 13 m tall, four-year-old, plantation Eucalyptus globulus Labill. in southern Tasmania. Responses were measured in three crown zones (lower, middle, upper) over a period of 11 months. Height increment was unaffected by defoliation, but diameter increment was significantly reduced 155 days after treatment. Defoliation treatment had no effect on stem volume and biomass partitioning compared with the control treatment. Trees responded to defoliation by decreased branch senescence in the lower crown, greater leaf area development in the mid crown and increased specific leaf area. Defoliation reduced concentration of soluble sugars (SS) in foliage by 22% and the pools of SS in the coarse roots by 34%. Decrease in root SS was only observed in 10-15 mm diameter class and the rootball. We concluded that this four-year-old E. globulus stands with a closed canopy was able to tolerate a single, partial artificial defoliation event, which is similarly observed with younger trees.     

Belowground competition from overstory trees influences Douglas-fir sapling morphology in thinned stands

We evaluated effects of belowground competition on morphology of naturally established coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) saplings in 60- to 80-year-old thinned Douglas-fir stands in southwestern Washington. We separately quantified belowground competition from overstory and understory sources using trenching and understory removal. In this light-limited environment of 26 ± 16% (std. dev.) full sunlight, 2-year exclusion of tree root competition by trenching increased sapling stem biomass by 18%, total aboveground biomass by 21%, number of interwhorl buds by 68%, total foliar biomass by 33%, and foliar biomass on branch components over 4 years old by 143%. Belowground competition did not influence shoot:root ratio or foliar efficiency (i.e., stem growth per unit foliage biomass). Sapling needle size, specific leaf area, and internodal distance also were not affected by belowground competition; these variables were apparently a function of the low-light environment. The principal source of belowground competition was roots of overstory trees; effects of belowground competition from understory vegetation were minor. Thus, under a partial overstory, morphology of Douglas-fir regeneration was influenced by both belowground and aboveground competition from overstory trees. In this environment, understory vegetation control would not likely influence belowground competition to an extent that would affect sapling morphology.     

The effects of acid rain and ozone on biomass and leaf area parameters of shortleaf pine (Pinus echinata Mill.)

Shortleaf pine (Pinus echinata Mill.) seedlings in 24 open-top chambers were exposed to combinations of ozone (carbon-filtered (control), ambient, 1.7 x ambient, and 2.5 x ambient) and acidic precipitation (pH 5.3, 4.3 and 3.3) for 16 months (1989 harvest) or 28 months (1990 harvest). Although the effects of acid rain were generally not significant, there was a trend toward increased aboveground biomass and leaf area in seedlings subjected to the low pH treatments. Because N concentrations in the soils generally increased with decreasing pH, we concluded that the effects of acid rain on aboveground biomass and leaf area were a consequence of an increasing concentration of soil N. In the 1989 harvest, seedlings in the 2.5 x ambient ozone treatment had significantly less biomass in all aboveground plant components and significantly less total leaf area than seedlings in the 1.7 x ambient ozone treatment. In the 1990 harvest, there were no significant effects of ozone on total aboveground biomass, although there was a trend toward reduced biomass in seedlings in the 2.5 x ambient ozone treatment. Both total leaf area and leaf biomass were significantly less in seedlings exposed to 2.5 x ambient ozone for 28 months than in both control seedlings and seedlings in the 1.7 x ambient ozone treatment. The greater, but not always significant, aboveground biomass and leaf area of seedlings in the 1.7 x ambient ozone treatment compared with control seedlings may be associated with the observed increase in soil nitrate concentration as a result of increased rates of leaf senescence and litterfall.     

Evaluating a simple radiation/dry matter conversion model using data from Eucalyptus globulus plantations in Western Australia

A simple model that describes growth in terms of physical and physiological processes is needed to predict growth rates and hence the productivity of trees at particular sites. The linear relationship expected between absorbed photosynthetically active radiation (phi(pa), MJ m(-2)) and dry mass production (G(t)); i.e., G(t) = epsilonphi(pa), where epsilon is the radiation utilization coefficient, was fitted to three years' data from five Western Australian Eucalyptus globulus Labill. plantations for which monthly growth measurements, leaf area indices, weather data and soil water measurements were available. Reductions in growth efficiency relative to absorbed photosynthetically active radiation were associated with high vapor pressure deficits (D, kPa) so the relationship between monthly aboveground biomass increments and D was used to calculate utilizable phi(pa). Plotting cumulative aboveground growth against utilizable phi(pa) gave strong linear relationships with slope epsilon. Values of epsilon ranged from 0.93 to 2.23 g dry mass MJ(-1) phi(pa). The variation could not be explained either in terms of soil water content in the root zones, because all plantations appeared to have access to groundwater, or in terms of soil chemistry. A value of epsilon approximately 2.2 is considered near the maximum likely to be applicable to Eucalyptus plantations. An interesting peripheral finding was a strong relationship between allometric ratios and soil phosphorus; this, if confirmed elsewhere, will be of considerable value in converting biomass increments to wood production. There was also a strong negative relationship between the average ratio of leaf/total aboveground biomass and soil nitrogen content.     

Mechanical perturbation affects conductivity, mechanical properties and aboveground biomass of hybrid poplars

Xylem development in trees is affected by dynamic mechanical stresses imposed on stems by wind. To assess clonal differences in response to mechanical perturbation (MP), we subjected seven greenhouse-grown F1 hybrids of Populus trichocarpa Torr. and A. Gray. x P. deltoides Bartr. ex Marsh. to a standard MP treatment consisting of 20 manually imposed stem flexures per day for 70-90 days. Effects of MP on aboveground biomass, hydraulic conductivity (k(h)), specific conductivity (k(s)), flexural stiffness (EI), modulus of elasticity (MOE) and modulus of rupture (MOR) were determined. Treatment increased stem radial growth and decreased height growth, leaf area and total aboveground biomass. It also significantly decreased k(s), MOE and MOR, but significantly increased EI and wood specific gravity in most clones. Mechanical perturbation caused greater stem rigidity, without having a significant effect on whole-stem k(h) or percent loss of conductivity due to embolism. Maximum k(h) was positively correlated with EI in both control (r(2) = 0.54, P < 0.0001) and MP-treated (r(2) = 0.61, P < 0.0001) plants, and k(s) and MOE were positively correlated with percent vessel lumen area (r(2) = 0.45, P < 0.0001 and r(2) = 0.28, P = 0.002, respectively). Thus, contrary to our expectation of a trade-off between conductivity and wood strength, there may be an opportunity to select clones for woody biomass production that are superior in both mechanical strength and hydraulic conductivity, as is the triploid Clone 19-61.     

Quantitative relationships between fine roots and stand characteristics

Fine roots absorb nutrients and water for photosynthesizing leaves, which in return provide them with hydrocarbon products. Knowledge of the fine root biomass (FRB) at the individual tree level and its relationships with other components related to tree growth, especially leaves aboveground, is scarce. Therefore, we reviewed the FRB of major forest-forming species using a database of 518 forest stands compiled from the literature, including 21 tree species and 16 shrub species, in order to confirm the relationships between environmental or forest stand variables and FRB at the stand and tree levels, and we further determine the relationships between fine roots belowground and leaves aboveground. Correlations between FRB and site characteristics (latitude, elevation, age, density, and basal area) appeared to be species-specific. There were hardly any significant correlations between stand FRB and latitude, elevation, stand age and stand density. Tree FRB was better correlated with tree basal area than stand FRB with stand basal area. There was a significant linear relationship between tree FRB and tree basal area. In addition, individual FRB was significantly linearly related to leaf biomass for all analyzed species. When these species were grouped into coniferous and deciduous, or all species together, there were still significant linear relationships between tree FRB and tree basal area and leaf biomass. The ratios of FRB to leaf biomass varied between and among species and even among regions for the same species. For both Picea abies and Pinus sylvestris, the ratio of FRB to leaf biomass was negatively related to the ratio of annual actual evapotranspiration to annual potential evapotranspiration, which was an indicator of water availability.     

Light distribution,photosynthetic rate and yield in a Paulownia-wheat intercropping system in China

Measurements of photosynthetically active radiation (PAR), leaf photosynthesis, canopy leaf area index (LAI) and crop yield were carried from flowering to maturity to study the effects of tree shading on crop yield in a Paulownia-wheat intercropping system in China. We found that the tree shading reduced the amount of incoming PAR within the intercropping system by 22%, 44% and 56% during flowering, grain-filling and maturing, respectively. The amount of PAR intercepted by the wheat crop inside the intercropping system was 34%, 55% and 68% less than the crop outside the intercropping system during flowering, grain-filling and maturing. Estimates of PAR-saturated leaf photosynthetic rate were not affected by tree shading, and the differences between the wheat yield inside the intercropping system and outside the intercropping system can be explained by the difference in the amount of PAR intercepted. Total grain numbers and grain dry weight per 1000 grains were linearly correlated with the amount of PAR intercepted during 7 days prior to anthesis and during anthesis and grain-filling, respectively. As compared with the wheat crop outside the intercropping system, grain numbers and grain dry weight were reduced by 36% and 25%, respectively. As a result wheat yield inside the intercropping system was 51% lower than that outside the intercropping system. Our results suggest a much higher reduction in crop yield and much smaller economic gain of the Paulownia-wheat intercropping system than previous studies on the similar Paulownia-wheat intercropping systems in China.     

Planting spacing affects canopy structure,biomass production and stem roundness in poplar plantations

To improve the productivity and wood quality of poplar plantations, effects of four planting spacing on canopy characteristics, biomass production and stem roundness in poplar plantations were evaluated over 8 years. Planting spacing influenced canopy characteristics of the plantations, and further affected the understory vegetation and plantation productivity. Understory vegetation biomass and Shannon-Wiener index were negatively correlated with leaf area index, but both diversity indexes and aboveground biomass of understory vegetation were higher in stands with a wider spacing. Tree diameter growth increased with increasing planting spacing, while the increment in plantations of square configurations (5?×?5 m and 6?×?6 m) was higher than those with rectangular configurations (3?×?8 m, 4.5?×?8 m). The highest poplar biomass production was achieved in the plantation with 5?×?5 m spacing at age 8. Moreover, poplar trees showed a tendency with better stem roundness in a square configuration. The results suggest that planting spacing not only affect canopy characteristics, understory vegetation and tree growth but also wood quality, and square configurations (5?×?5 m and 6?×?6 m) could be a better option for poplar plywood timber production at similar sites.     

Allometric relations for biomass partitioning of Nothofagus antarctica trees of different crown classes over a site quality gradient

Data on tree biomass are essential for understanding the forest carbon cycle and plant adaptations to the environment. We determined biomass accumulation and allometric relationships in the partitioning of biomass between aboveground woody biomass, leaves and roots in Nothofagus antarctica. We measured above- and belowground biomass of N. antarctica trees across different ages (5–220 years) and crown classes (dominant, codominant, intermediate and suppressed) in three site qualities. The biomass allocation patterns were studied by fitting allometric functions in biomass partitioning between leaves (M_L), stem and branches (M_S) and roots (M_R). These patterns were tested for all pooled data and according to site quality and crown classes. Biomass accumulation varied with crown class and site quality. The root component represented 26–72% of the total biomass depending on age and site. N. antarctica scaling exponents for the relationships M_Lvs. M_S, M_Avs. M_R, and M_Svs. M_R were close to those predicted by the allometric biomass partitioning model. However, when biomass allocation was analyzed by site quality the scaling exponents varied following the optimal partitioning theory which states that plants should allocate more biomass to the part of the plant that acquires the most limiting resource. In contrast, the crown class effect on biomass partitioning was almost negligible. In conclusion, to obtain accurate estimations of biomass in N. antarctica trees the allometric approach appears as an useful tool but the site quality should be taken into consideration.     

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.     

Interactive effects of nitrogen and water availabilities on gas exchange and whole-plant carbon allocation in poplar

Cuttings of balsam spire hybrid poplar (Populus trichocarpa var. Hastata Henry x Populus balsamifera var. Michauxii (Dode) Farwell) were grown in sand culture and irrigated every 2 (W) or 10 (w) days with a solution containing either 3.0 (N) or 0.5 (n) mol nitrogen m(-3) for 90 days. Trees in the WN (control) and wn treatments had stable leaf nitrogen concentrations averaging 19.4 and 8.4 mg g(-1), respectively, over the course of the experiment. Trees in the Wn and wN treatments had a similar leaf nitrogen concentration, which increased from 12.0 to 15.8 mg g(-1) during the experiment. By the final harvest, mean stomatal conductances of trees in the wN and wn treatments were less than those of trees in the Wn and WN treatments (1.8 versus 4.6 mm s(-1)). Compared to the WN treatment, biomass at the final harvest was reduced by 61, 72 and 75% in the Wn, wN and wn treatments, respectively. At the final harvest, WN trees had a mean total leaf area of 4750 +/- 380 cm(2) tree(-1) and carried 164 +/- 8 leaves tree(-1) with a specific leaf area of 181 +/- 16 cm(2) g(-1), whereas Wn trees had a smaller mean total leaf area (1310 +/- 30 cm(2) tree(-1)), because of the production of fewer leaves (41 +/- 6) with a smaller specific leaf area (154 +/- 2 cm(2) g(-1)). A greater proportion of biomass was allocated to roots in Wn trees than in WN trees, but component nitrogen concentrations adjusted such that there was no Wn treatment effect on nitrogen allocation. Compared with WN trees, rates of photosynthesis and respiration per unit weight of tissue of Wn trees decreased by 28 and 31%, respectively, but the rate of photosynthesis per unit leaf nitrogen remained unaltered. The wN and Wn trees had similar leaf nitrogen concentrations; however, compared with the Wn treatment, the wN treatment decreased mean total leaf area (750 +/- 50 cm(2) tree(-1)), number of leaves per tree (29 +/- 2) and specific leaf area (140 +/- 6 cm(2) g(-1)), but increased the allocation of biomass and nitrogen to roots. Net photosynthetic rate per unit leaf nitrogen was 45% lower in the wN treatment than in the other treatments. Rates of net photosynthesis and respiration per unit weight of tissue were 48 and 33% less, respectively, in wN trees than in Wn trees.     

Estimates of forest biomass in the Brazilian Amazon: New allometric equations and adjustments to biomass from wood-volume inventories

Uncertainties in biomass estimates in Amazonian forests result in a broad range of possible magnitude for the emissions of carbon from deforestation and other land-use changes. This paper presents biomass equations developed from trees directly weighed in open forest on fertile soils in the southern Amazon (SA) and allometric equations for bole-volume estimates of trees in both dense and open forests. The equations were used to improve the commonly used biomass models based on large-scale wood-volume inventories carried out in Amazonian forest. The biomass estimates from the SA allometric equation indicate that equations developed in forests on infertile soils in central Amazonia (CA) result in overestimates if applied to trees in the open forests of SA. All aboveground components of 267 trees in open forests of SA were cut and weighed, and the proportion of the biomass stored in the crowns of trees in open forest was found to be higher than in dense forest. In the case of inventoried wood volume, corrections were applied for indentations and hollow trunks and it was determined that no adjustment is needed for the form factor used in the RadamBrasil volume formula. New values are suggested for use in models to convert wood volume to biomass estimates. A biomass map for Brazilian Amazonia was produced from 2702 plots inventoried by the RadamBrasil Project incorporating all corrections for wood density and wood volume and in factors used to add the bole volume of small trees and the crown biomass. Considering all adjustments, the biomass map indicates total biomass of 123.1 Gt (1 Gt = 1 billion tons) dry weight (aboveground + belowground) for originally forested areas in 1976 in the Brazilian Legal Amazon as a whole (102.3 Gt for aboveground only) at the time of the RadamBrasil inventories, which were carried out before intensive deforestation had occurred in the region. Excluded from this estimate are 529,000 km² of forest lacking sufficient RadamBrasil inventory data. After forest losses of 676,000 km² by 2006 – not counting 175,000 km² of this deforested area lacking RadamBrasil data – the estimated dry biomass stock was reduced to 105.4 and 87.6 Gt (aboveground + belowground and only above-ground). Thus, in 2006 the carbon storage in forested areas in Brazilian Amazonia as a whole will be around 51.1 Gt (assuming 1 Mg dry biomass = 0.485 Mg C). Biomass estimates by forest type (aggregated into 12 vegetation classes) are provided for each state in the Brazilian Legal Amazon.     

Increasing wood production through old age in tall trees

How long forest trees can sustain wood production with increasing age remains an open question, primarily because whole-crown structure and growth cannot be readily measured from the ground or on felled trees. We climbed and directly measured crown structures and growth rates of 43 un-suppressed individuals (site trees) of the two tallest species – Eucalyptus regnans and Sequoia sempervirens – representing a wide range of tree sizes and ages. In both species, ground-level measurements of annual growth, including height, ring width, and basal area increment, exhibited the oft-reported trend of decreasing growth (or no change in growth) with age, yet wood production of the entire main trunk and whole crown both increased with size and age up to and including the largest and oldest trees we measured. The balance between structural metrics of whole-crown respiratory demands (cambium area, inner bark volume, sapwood volume, and heartwood deposition area) and photosynthetic capacity (leaf area and green bark area) was statistically independent of size but not age. After accounting for the effect of size, trees with lower potential respiratory demands grew more than trees with higher potential respiratory demands per unit photosynthetic area. The strongest determinant of tree energy balance was the ratio of aboveground cambium area to leaf area. Among the site trees we examined, over 85% of the variation in annual wood production was explained by variation in size, and the proportion of total aboveground wood production in appendages (branches, limbs, and reiterated trunks) increased linearly with size. With increasing age in both species, the proportion of annual wood production converted to heartwood increased in main trunks and appendages. The oldest tree we measured produced more heartwood in its main trunk over 651 years (351 m³) than contained in any tree we measured <1500 years old. The two tallest tree species achieve similar stature despite divergent growth dynamics and ecologies. At one extreme, E. regnans attains great size quickly but dies relatively young because trees are susceptible to fire and fungi. At the other extreme, S. sempervirens attains great size more slowly but has a long lifespan because trees resist fire and prioritize investment in decay-resistant heartwood. Increasing wood production as trees age is a mechanism underlying the maintenance of biomass accumulation during forest development and the carbon-sink capacity of old-growth forests.     

The effect of red wood ant (<Emphasis Type="Italic">Formica rufa</Emphasis> group) mounds on root biomass,density, and nutrient concentrations in boreal managed forests

Red wood ants (Formica rufa group, RWAs) are common insects in boreal forests in Fennoscandia, and they build large, long-lived mounds as their nests. RWA mounds are enriched with carbon and nutrients, but little information is available about how they affect root distribution and the nutrient uptake of trees. In this study, we investigated the biomass, biomass density, nutrient concentrations, and amounts of fine (<2 mm) and coarse (>2 mm) roots in RWA mounds, and compared them with those of surrounding forest soil in mixed coniferous stands of different age classes in Finland. Neither fine nor coarse root biomasses differed significantly between the aboveground parts of the mounds and the organic layer of the soil. Root biomass density was lower in mounds than in the organic layer. However, fine root biomass and biomass density were higher in the belowground parts of mounds than in the surrounding mineral soil. Macroelement (N, Ca, K, P, S, Mg) and Zn and Cu concentrations in roots in the mounds were significantly higher than those in the organic layer. Root biomass and biomass density did not differ between stands of different age classes. The results of this study indicate that RWA mounds increase heterogeneity in root distribution in forest ecosystems, and also increase the availability of nutrients for plants that extend their roots inside RWA mounds.