Crown architecture and stemwood production in Norway spruce (Picea abies (L.) Karst.)

The morphological data for Norway spruce (Picea abies (L.) Karst.) trees published by Burger (1953) were reanalyzed to examine the relationships between crown shape, needle and branch mass density of crown volume, needle efficiency in stemwood production, harvest index and stemwood production efficiency per unit of crown projected area. Production of stemwood per unit of crown projected area was higher in narrow-crowned trees than in broad-crowned trees because the narrow-crowned trees had (i) smaller horizontal space requirements for equal crown volumes, (ii) higher leaf area index due to both the geometrical crown shape and higher needle density per unit of crown volume, and (iii) higher harvest index indicating high allocation of dry matter production to stem.     

A model for the relationship between branch number and biomass in Pinus sylvestris crowns and the effect of crown shape and stand density on branch and stem biomass

The branch biomass of young Scots pines (Pinus sylvestris L.) was inversely related to the branch number per projected crown area giving greater branch density and smaller branch biomass for narrow crowns than for broad crowns. In particular, in dense stands the small share of branches from the total biomass of narrow‐crowned trees was emphasized. The productivity of narrow crowned trees was, however, smaller than that of trees with broad crowns. The concept of the tree ideotype has been discussed on the basis of theoretical calculations.     

辽东山区日本落叶松生物量相容性模型的研究

基于60株辽东山区日本落叶松样木生物量的实测数据,分析不同林龄条件下立木各部分生物量的变化情况,并应用度量误差方法建立立木相容性生物量模型。结果表明:树叶、树枝、树皮生物量占总生物量的比值随林龄增长呈下降趋势,干材占总生物量的比值随林龄增长呈上升趋势。在筛选出总生物量与各分量最优独立模型的基础上,应用三级控制的方法建立生物量相容性模型,并采用加权回归方法消除总量和各分量模型的异方差。建立的总量、地上部分、树干、干材、树皮生物量模型,其R2均大于0.9;树根、树冠、树叶和树枝生物量的R2略低,介于0.7 0.9之间。通过独立样本对模型的相容性和预测精度进行检验,各分量预测值所占总生物量的百分比之和为1,模型完全相容;根、冠、叶和枝的模型预测精度略低于90%,其他部位模型的预测精度都在95%以上,模型的预测精度较高。     

Inherited narrow crown form, harvest index and stem biomass production in Norway spruce, Picea abies

The effect of crown form on stem biomass production was investigated in an 18-19-year-old Norway spruce stand (Picea abies (L.) Karst.). The harvest index was 0.271 in pendula trees, which have a heritable narrow crown form, and 0.235 in normal-crowned trees and the dry weights of stem biomass were 2.57 kg and 3.37 kg, respectively. However, the production of stem biomass per crown projected area was more than twice as much in pendula trees as in normal-crowned trees. Results indicate that the crown form of pendula spruce is less plastic than that of normal-crowned spruce suggesting that such trees might maintain a high stemwood yield per unit ground area with increasing stand density. Because pendula spruce has a higher harvest index and a less flexible crown than normal-crowned spruce, it might be useful for crop tree ideotype breeding.     

Vegetation competition effects on aboveground biomass and macronutrients,leaf area,and crown structure in 5-year old Douglas-fir

Vegetation control (VC) in forest plantations often increases growth of crop trees but can also affect biomass and nutrient partitioning to tree components. We examined above-ground biomass and macronutrients, leaf area and crown structure in 5-year old Douglas-fir (Pseudotsuga menziessi (Mirb.) Franco) growing with VC and with no vegetation control (NVC) in coastal Washington, United States of America. Trees in VC had larger stem, branch, foliage and total biomass than trees of equal stem diameter at 1.3 m above ground in NVC. The difference in component biomass between treatments was in the order: branch > foliage > stem. Trees in the VC regime did not differ in macronutrient concentration in stem, branch and foliage except for branch N which was greater in NVC than in VC. Differences in tree macronutrient stores between VC and NVC ranged from 2.2 times for Mg to 2.6 times for N and K. The relationship between stem diameter and leaf area was linear in both VC treatments. The relationship between stem diameter and foliage biomass was curvilinear for both VC treatments. Results of this study support the need for separated allometric equations to estimate biomass for young Douglas-fir growing in areas with and without VC.     

Patterns in aboveground carbon allocation and tree architecture that favor stem growth in young Scots pine from high latitudes

The production and allocation of aboveground biomass and the characteristics of tree architecture were examined in eight-year-old Scots pine (Pinus sylvestris L.). Considerable among-tree variation existed in tree architecture, total aboveground dry mass production, and dry mass partitioning among tree parts. A linear relationship existed between needle and branch mass. Stem mass was directly proportional to tree height, which in turn was directly proportional to the allocation ratio between stem mass and total needle + branch mass production. The architectural characteristics that were related to a high proportional allocation to stem and high stemwood production were a large mean shoot volume, large mean number of branches per whorl, long needle retention and a high crown length/crown width ratio. Individual trees were found that combined high stemwood production with both high harvest index and high stemwood specific gravity.     

Variation in the ratio of shoot silhouette area to needle area in fertilized and unfertilized Norway spruce trees

We compared the range and variation in shoot silhouette area to projected leaf area ratio (SPAR) in fertilized and unfertilized (control) Norway spruce (Picea abies (L.) Karst.) trees. We measured SPAR for several view directions of 169 shoots at different depths in the crown of fertilized and control trees. There was an increase in SPAR with depth in the crown in both control and fertilized trees. In the fertilized trees, however, mean SPAR was larger overall, the increase with depth in the crown was steeper, and there was a larger variation in SPAR with inclination and rotation angle of the shoot (relative to the view direction). In particular, shoots in the lower crown of fertilized trees were rotationally asymmetrical ("flat") and had high values of the maximum ratio of shoot silhouette area to projected leaf area (SPAR(max)). Differences in SPAR between fertilized and control trees were explained by changes in shoot structure in response to fertilization and shading. Shoots of fertilized trees were larger and had more needle area than shoots of control trees. However, the ratio of needle area to shoot size was smaller in fertilized trees than in control trees, implying less within-shoot shading and, consequently, a larger SPAR. Also, the increase in SPAR with increased shading (depth in the crown) could be explained by a decrease in the ratio of needle area to shoot size. In addition, because fertilized trees had more needle area than control trees, the effect of shading at a given depth in the crown was more pronounced in fertilized trees than in control trees.     

Carbon concentrations of tree components,forest floor and understorey in young Pinus sylvestris stands in north-western Turkey

Abstract

This study examines the carbon concentrations of various above-ground biomass components (stem, bark, living branch, dead branch, needle and above-ground biomass), the forest floor and understorey. The research was carried out on Scots pine (Pinus sylvestris L.) stands in north-western Turkey. Sample trees were cut, and forest floor and understorey samples were collected from the 13 sampling areas in the 31-year-old Scots pine stands. Carbon contents of these samples were determined. Carbon concentrations of all tree components were found to be higher than 50%. This value is referred to as the general carbon conversion factor (CF). The weighted mean carbon concentration was found to be 51.96%. Consequently, if the CF for a single tree is taken to be 50%, carbon storage in the above-ground parts of the tree would have been underestimated by 3.77%. Carbon concentration, which was 53.02% in needles, was reduced to 50.08% in the litter plus fermentation layer of the forest floor and to 40.08% in the humus layer owing to decomposition. In the understorey, carbon concentration was determined to be 47.64%. There was a significant difference between the carbon concentrations of the tree components, forest floor and understorey.     

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.     

Crown characteristics of juvenile loblolly pine 6 years after application of thinning and fertilization

Total foliage dry mass and leaf area at the canopy hierarchical level of needle, shoot, branch and crown were measured in 48 trees harvested from a 14-year-old loblolly pine (Pinus taeda L.) plantation, six growing seasons after thinning and fertilization treatments.

In the unthinned treatment, upper crown needles were heavier and had more leaf area than lower crown needles. Branch- and crown-level leaf area of the thinned trees increased 91 and 109%, respectively, and whole-crown foliage biomass doubled. The increased crown leaf area was a result of more live branches and foliated shoots and larger branch sizes in the thinned treatment. Branch leaf area increased with increasing crown depth from the top to the mid-crown and decreased towards the base of the crown. Thinning stimulated foliage growth chiefly in the lower crown. At the same crown depth in the lower crown, branch leaf area was greater in the thinned treatment than in the unthinned treatment. Maximum leaf area per branch was located nearly 3–4 m below the top of the crown in the unthinned treatment and 4–5 m in the thinned treatment. Leaf area of the thinned-treatment trees increased 70% in the upper crown and 130% in the lower crown. Fertilization enhanced needle size and leaf area in the upper crown, but had no effect on leaf area and other variables at the shoot, branch and crown level. We conclude that the thinning-induced increase in light penetration within the canopy leads to increased branch size and crown leaf area. However, the branch and crown attributes have little response to fertilization and its interaction with thinning.     

Branch mortality and potential litterfall from Douglas-fir trees in stands of varying density

Forest floor characteristics influence nutrient cycling and energy flow properties of forest ecosystems, and determine quality of habitat for many forest plants and animals. Differential crown recession and crown development among stands of differing density suggest that an opportunity may exist to control the input of fine woody litter into the system by manipulating stand density. The objective was to measure the rate of branch mortality among stands of differing density and to estimate the range in total per hectare necromass inputs. Although litter traps are reliable for estimating per hectare rates of litterfall, branch mortality dating on sectioned stems uniquely allows assessment of several other litterfall components: (1) individual tree contributions to total litterfall; (2) the amount of branch material released by mortality, regardless of whether the branches are shed to the forest floor; (3) the distribution of basal diameters characterizing the litterfall from a given tree and stand. Twenty-four trees were felled and sectioned on permanent plots that were part of a silvicultural study of stand density regimes in Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco.). Whorl branches were dissected out of bole sections to determine the dates of mortality, and a branch biomass equation was applied to estimate potential rate of litterfall. Periodic annual rates were expressed in four ways: (1) number of branches per tree; (2) mass of branches per tree; (3) mass of branches per unit of crown projection area; (4) mass of branches per hectare. For the growth periods investigated, larger trees and trees growing on denser plots tended to release a greater necromass through branch mortality. Average branch basal diameter generally decreased with increasing stand density. Annual branch mortality ranged from 33 to 430 g m⁻² crown projection area for individual trees, and from 236 to 1035 kg ha⁻¹ for individual plots. These rates approached the low end of the range of previously published fine litterfall rates for Douglas-fir. Rates on these plots were relatively low owing to the temporary delay in crown recession imposed by artificial thinning. A conceptual model of branch litter dynamics is presented to depict consistencies with crown development among stands managed under different density regimes.     

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.     

Crown architecture of Abies balsamea from four canopy positions

Data collected from four distinct canopy positions from each of 39 Abies balsamea (L.) Miller trees were used to construct models to describe the cumulative leaf area distribution within the crown and to predict the needle mass of individual branches, the average branch angle, branch diameter, branch length, and crown radius per whorl, and the average number of living branches per whorl. We tested the hypotheses that regression models are equal among canopy positions and that a model to predict branch needle mass is valid at the northern and southern extremes of the central climatic zone of Maine. Canopy position had an effect on the models constructed to predict needle mass, branch angle, branch diameter, branch length, crown radius, and the number of living branches per whorl. However, compared with an expanded model that incorporated parameters calculated for each crown class, there was only a small loss in model precision when a general model constructed from data pooled from all crown classes was used to predict needle mass, branch angle, and branch diameter. Regression equations unique to each crown class were needed to predict crown shape and leaf area distribution in the crown satisfactorily. Our branch needle mass model, which was constructed from data collected at the southern extreme of the central climatic zone of Maine, consistently underestimated needle branch mass when applied to the northern extreme of the central climatic zone.     

Examining age- and altitude-related variation in tree architecture and needle efficiency in Norway spruce using trend surface analysis

The aim of this study was to examine the variation in tree architecture and needle efficiency in stemwood production in Norway spruce in relation to tree age and altitude of growing site. The data, which were obtained from the literature, described individual Norway spruce (Picea abies (L.) Karst.) trees from even-aged stands in Switzerland. Second-order trend surface models, with tree age and altitude as independent variables, were used in the analysis. The fitted models for stem, branch and needle dry masses explained 95%, 75% and 64% of the variation, respectively. The model for the estimated mean branch density in the crown (kg m⁻³) explained 64% of the variation and the model for mean needle density in the crown (kg m⁻³) only 28% of the variation. Crown structural characteristics, which showed age- and altitude-related variations, included live crown ratio (59% of variation explained), number of living whorls (43% of variation explained), mean weight of single needle (40% of variation explained) and specific needle area (27% of variation explained). Tree age had a strong effect on needle efficiency in stemwood production, so that needle efficiency increased up to the age of 50–70 years, depending on altitude.     

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.     

Crown structure and leaf area index development in thinned and unthinned Eucalyptus nitens plantations

The crown structure of Eucalyptus nitens (Deane & Maiden) Maiden 6 years after thinning, and the development of stand leaf area index both immediately and 6 years after thinning, were investigated. Thinning did not alter branch angle, branching density or the relationship between branch size and branch leaf area. However, larger branches were found in the lower crown of thinned trees and the increase in leaf area as a result of thinning occurred on the northern aspect of the crown. The vertical distribution of leaf area in unthinned trees was skewed toward the top of the crown and correlated with live crown ratio. The vertical distribution of leaf area in thinned trees tended to be less skewed and was unrelated to tree size or dominance. Leaf area index, as estimated from light interception measurements, increased at a constant rate soon after thinning regardless of residual stocking. In the longer term, residual stocking had a strong influence on leaf area increase per tree and was correlated with changes in crown length.     

Crown architecture of stand-grown sugar maple (Acer saccharum Marsh.) in the Adirondack Mountains

Leaf and crown morphology of shade-tolerant sugar maple (Acer saccharum Marsh.) were examined to test the hypotheses (1) that leaf area exhibits significant plasticity both within and between crown classes and individual tree crowns and (2) that leaf area is accurately predicted from estimates of crown volume. A total of 18 trees, ranging from 3.3 to 43.4 cm dbh, were felled and dissected into upper, middle, lower, and below-crown layers, for measurements of leaf, bark, and xylem dimensions. For dominant trees only, bark thickness and xylem radii were higher within the crown than below the crown. Cumulative leaf area index increased with decreasing stratum height at similar rates in all trees, except for two trees that were located in the understory. Area leaf weight declined with decreasing stratum height within the crown of all except four overstory trees. These four trees showed an increase with decreasing stratum height, i.e., leaves were heavier per unit area in the lower crown stratum and below the crown than they were at mid-crown. Within-tree leaf area density was usually higher in the upper crown of overstory trees and in the lower crown of understory trees. Total crown volume was the best predictor of whole-tree leaf area, but it was only slightly better than dbh.     

Changes in physiological attributes of ponderosa pine from seedling to mature tree

Plant physiological models are generally parameterized from many different sources of data, including chamber experiments and plantations, from seedlings to mature trees. We obtained a comprehensive data set for a natural stand of ponderosa pine (Pinus ponderosa Laws.) and used these data to parameterize the physiologically based model, TREGRO. Representative trees of each of five tree age classes were selected based on population means of morphological, physiological, and nearest neighbor attributes. Differences in key physiological attributes (gas exchange, needle chemistry, elongation growth, needle retention) among the tree age classes were tested. Whole-tree biomass and allocation were determined for seedlings, saplings, and pole-sized trees. Seasonal maxima and minima of gas exchange were similar across all tree age classes. Seasonal minima and a shift to more efficient water use were reached one month earlier in seedlings than in older trees because of decreased soil water availability in the rooting zone of the seedlings. However, carbon isotopic discrimination of needle cellulose indicated increased water-use efficiency with increasing tree age. Seedlings had the lowest needle and branch elongation biomass growth. The amount of needle elongation growth was highest for mature trees and amount of branch elongation growth was highest for saplings. Seedlings had the highest biomass allocation to roots, saplings had the highest allocation to foliage, and pole-sized trees had the highest allocation to woody tissues. Seedlings differed significantly from pole-sized and older trees in most of the physiological traits tested. Predicted changes in biomass with tree age, simulated with the model TREGRO, closely matched those of trees in a natural stand to 30 years of age.     

Biomass,stem basic density and expansion factor functions for five exotic conifers grown in Denmark

Adequate allometric equations are needed for estimating carbon pools of fast growing tree species in relation to international reporting of CO₂ emissions and for assessing their possible contribution to increasing forest biomass resources. We developed models for predicting biomass, stem basic density and expansion factors of stem to above-ground biomass for five fast growing conifers. Data included destructive measurements of 236 trees from 14 sites, covering a wide range of growth conditions. To ensure model efficiency, models for predicting stem, crown and total above-ground biomass for the five species were estimated simultaneously using a linear, mixed effects model that allowed contemporaneous correlations between the different tree components. Models differed among species and included dbh and tree height. The models explained more than 98% of the variation in above-ground biomass and reflected differences in the allometry between tree species. Stem density differed among species but generally declined with increasing site index and dbh. The overall model for predicting stem basic density included dbh, H₁₀₀ and site index and explained 66% of the total variation. Expansion factors decreased from 1.8–2.0 in small trees (dbh < 10 cm) to 1.1–1.2 for large trees (dbh > 25 cm), but differed among species. The overall model explained 86% of the variation and included quadratic mean diameter and dbh.     

Pipe-model ratio distributions and branch foliage biomass: differences between two sympatric spruce species

The foliage biomass–sapwood relationship (the pipe model) is critical for tree growth and is used in tree growth models for understanding the implications of this structural relationship on the allocation of resources. In this research, we compared this relationship for two commercially important and sympatric species, black spruce (Picea mariana (Mill.) B.S.P.) and white spruce (Picea glauca (Moench) Voss). At locations in eastern Canada, 57 black and 50 white spruce trees were destructively sampled to obtain foliage biomass, crown structure, and tree stem measures. Using a model-based approach, we compared foliage biomass–branch basal area and foliage biomass–sapwood relationships at the tree and disk (i.e. along the tree stem) levels (i.e. pipe-model ratios) between these two species. We found that (i) branch foliage biomass–branch basal area was greater for black spruce than white spruce and (ii) pipe-model ratios along the tree stem given tree size were greater for black spruce than for white spruce. We attributed these differences to: (i) greater shade tolerance and leaf longevity of black spruce; (ii) slower growth rates of black spruce; and (iii) differing hydraulic strategies and mechanical requirements.