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.     

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.     

Genetic and environmental control of crown development in Picea sitchensis and its relation to stem wood production

Differences in stem wood production were found among four clones of Picea sitchensis (Bong.) Carr. at both an agricultural and a forest site. Clonal rankings were not consistent between the sites. Four variables were identified that may influence stem wood production, foliage amount, total aboveground production per unit foliage, proportional allocation of production to new foliage and to branch wood thickening. Comparison of clonal performance between sites showed that stem wood production could be influenced by differences in each of these variables. The four variables were themselves determined by components of crown structure, these were (i) production of new branches from the main stem, (ii) the numbers and lengths of branchlets supported on branches from the main stem, (iii) foliage production per unit branchlet length and foliage longevity, and (iv) branch wood thickening per unit branch length. These components varied both among clones and between sites. Branch production from the mainstem varied among clones, and for some clones it varied substantially between sites. In some clones branch production was positively correlated with mainstem height increment. There were differences among clones in the way that branchlet production varied between the agricultural and forest sites. In the absence of needle fall, foliage weight/branch length varied threefold among clones, but for each clone, varied little between sites. Branch wood weight/branch length was generally greater when foliage weight/branch length was large, but total branch wood increment/tree was strongly influenced by total branch length. The dynamics of crown development and its relation to stem wood production are discussed with reference to (i) the importance of needle longevity, (ii) the importance of tallness and narrowness of crowns and (iii) the importance of branch thickening, a process that competes with stem growth in the utilization of photosynthate. It is concluded that although each of these factors may be important, none singly accounts for the observed differences in stem wood production among genotypes and the way these change in response to the environment.     

Relationships between crown projected area and components of above-ground biomass in Norway spruce trees in even-aged stands: Empirical results and their interpretation

Sample tree material was reanalyzed in order to study the relationships between horizontal crown projected area and components of above-ground biomass in Norway spruce (Picea abies (L.) Karst.) trees growing in even-aged stands. The needle mass of dominant trees increased linearly with the increase in crown projected area, but in co-dominant and dominanted trees the increase in needle mass levelled off toward larger crown projected areas. The branch mass of dominant and co-dominant trees accumulated faster than linearly with increasing crown projected area, whereas in dominated trees an approximately linear relationship existed between these two variables. The increase in needle and branch mass per unit increase in crown projected area was highest in dominant trees and decreased to co-dominant and dominated trees, i.e. with tree position in the canopy. The stem mass accumulated obviously faster than linearly and similarly in all tree classes with the increase in crown projected area. The narrow crown shape indicated a high density of all components of above-ground biomass per unit of crown projected area.     

Aboveground biomass and nutrient allocation in an age-sequence of <Emphasis Type="Italic">Larix olgensis</Emphasis> plantations

Biomass and nutrient (N, P, K, Ca, Mg) stock in various aboveground tree components (stemwood, stembark, branches and leaves) were quantified in an age sequence of pure Larix olgensis planta- tions (20, 35, 53 and 69 years old) in Northeast China. The results show that the aboveground biomass allocation in various tree components was in the order of stemwood (62%-83%), branches (9%-21%), stembark (7%-11%) and leaves (1%-6%) for all stands. The proportion of stemwood biomass to total aboveground biomass increased whereas that of other tree components decreased consistently with stand age from 20 to 53 years old, but kept relatively constant with stand age from 53 and 69 years old. The nutrient allocation in various tree components generally followed the same pattern as the biomass allocation (i.e. stemwood > branches > stembark > leaves). The proportion of nutrient stock in leaves to total aboveground nutrient stock decreased consistently with increasing stand age, while that in stemwood increased with stand age from 20 to 53 years old but then decreased from 53 to 69 years old. The rate of nutrient removal for stands was estimated at different stand ages under different logging schemes, showing that the rate of nutrient removal would be unchanged when the rotation length was shortened to 20 years by the harvest of stem only, but greatly increased by the harvest of total aboveground biomass. The rate of nutrient removal would be a considerable reduction for all elements by debarking, especially for Ca.     

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.     

林木竞争对臭冷杉生物量分配的影响

用不同高度树干直径建立并比较臭冷杉各器官生物量方程,分析林木竞争对臭冷杉地上、地下生物量分配的影响。结果表明:臭冷杉不同高度树干直径中,胸径是预测各器官生物量的最可靠变量;利用不同高度树干直径建立各器官生物量方程均会高估小个体样木(直径≤10cm)的生物量,并且随着直径增大,预测误差也随之增大;臭冷杉地上生物量与地下生物量的比值(T/R)与树木年龄、单株生物量、整株生物量年均生长率及树高年均生长率间均没有显著相关性(P>0.05);随着竞争增强,臭冷杉树干生物量占单株生物量的比例逐渐减小,枝叶生物量比例逐渐增大,而粗根生物量比例则基本保持不变;胸径年均生长率、树高年均生长率及单株生物量年均生长率均随着竞争强度增大逐渐减小,而T/R值并不受林木竞争的影响。     

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.     

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.     

Functions for estimating aboveground biomass of birch in Norway

A suite of regional allometric aboveground biomass functions were derived for Betula pubescens and Betula pendula for Norwegian conditions. The data consisted of 67 trees sampled throughout Norway. A total of 14 component functions were developed for total aboveground, total stem, stemwood, stem bark, live crown, live branch, leaf, and dead branch biomass using combinations of diameter at breast height and height as predictor variables. Application of the derived functions to existing local southern Norwegian mountain birch and regional Swedish biomass datasets indicated an overall good predictive ability of the developed functions. However, the functions produced slight underestimates, suggesting that the respective birch populations had differing biomass allocation patterns. When the developed functions were applied to Norwegian National Forest Inventory data, they produced slightly higher biomass stock and stock change estimates than what is obtained using existing Swedish functions. The higher estimates were evident in the north, central, and western part of Norway, while estimates were similar in southeastern Norway where growing conditions are most similar to Swedish conditions. The analysis indicates that the derived functions are the best available for regional birch biomass stock and stock change estimation in Norway.     

Morphological acclimation to understory environments in Abies amabilis, a shade- and snow-tolerant conifer species of the Cascade Mountains, Washington, USA

Light-related plasticity in a variety of crown morphology and within-tree characteristics was examined in sun and shade saplings of Abies amabilis Dougl. ex J. Forbes growing in two late-successional forests with different snow regimes in the Cascade Mountains of Washington, USA. Compared with sun saplings, shade saplings typically had broad flat crowns as a result of acclimation at several scales (needle, shoot, branch, crown and whole sapling). Shoots of shade saplings had a smaller needle mass per unit of stem length than shoots of sun saplings, a feature that enhances light-interception efficiency by reducing among-needle shading. The low annual rate of needle production by shade saplings was associated with a longer needle lifespan and slower needle turnover. Reduced needle production within a shoot was reflected at the branch level, with lateral branches of shade saplings having a smaller needle mass than branches of the same length of sun saplings. Reduced allocation to needles permits greater investment in branches and stems, which is necessary to support the horizontally expanding branch system characteristic of shade saplings. Mean branch age of shade saplings was significantly higher than that of sun saplings. Shade saplings had lower needle mass per unit of trunk biomass or total biomass, reflecting greater investment in the trunk as a support organ. Increased investment in support organs in shade was more evident in the snowier habitat. The observed morphological acclimation makes A. amabilis highly shade and snow-tolerant and thus able to dominate in many late-successional forests in snowy coastal mountain regions.     

Dependence of the aboveground respiration of hinoki cypress (Chamaecyparis obtusa) on tree size

Nighttime respiration was measured at monthly intervals over one year on the aboveground parts of five sample trees in an 8-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) stand, by an enclosed standing-tree method. The respiration rate rose rapidly from early spring to a maximum in June, and decreased abruptly in July and then gradually toward autumn and winter. The seasonal change in the respiration rate was synchronized with stem volume increment rather than with monthly mean air temperature. The respiration rate, r, of individual trees increased with increasing tree dimensions, such as stem volume, v(S), and stem girth at the base of the live crown, G(B). The dependence of respiration rate on tree size was successfully represented by a power function. The r - v(S) dependence was rather stronger than the r - G(B) (2) dependence, especially toward the end of the growing season (from July to September). The observed respiration rate was almost the same as the respiration rate corrected for the monthly mean air temperature. The annual respiration of individual trees was directly proportional to their phytomass or to its increment. Although the annual respiration of individual trees decreased proportionally to the square root of the leaf mass, it decreased abruptly in the range close to the smallest sample tree. Combining the monthly relationship between respiration rate and stem volume with the tree size distribution in the stand, the stand aboveground annual respiration was estimated to be 20.4 Mg CO(2) ha(-1) year(-1) (= 12.5 Mg dry mass ha(-1) year(-1)) for an aboveground biomass of 17.4 Mg ha(-1) with an annual increment of 6.51 Mg ha(-1) year(-1), i.e., the stand aboveground annual respiration amounted to the equivalent of 72% of the biomass or to almost twice the biomass increment.     

Improving models of forest nutrient export with equations that predict the nutrient concentration of tree compartments

? The objective of this study was to explore the distribution of major nutrients (N, P, K, Ca and Mg) in the aboveground compartments of an intensively managed tree species (Pinus pinaster Ait.). A total of 53 trees were cut down in even-aged stands respectively 8, 16, 26, 32 and 40 years old. The nutrient concentrations of the aboveground compartments were analysed.

? Nutrient concentrations of foliage did not vary with any of the variables used, except needle age. Nutrient concentrations of living branches, stem bark, stem sapwood, stem heartwood, stemwood and stem decreased with increasing branch diameter, bark thickness, sapwood thickness and heartwood thickness, respectively. Beyond a certain value of the predictive variable (stem diameter ≈ 15 cm; branch diameter ≈ 2.5 cm), the concentration of all the nutrients stabilised.

? A 50 year-old pine stand was used to obtain a validation dataset for nitrogen concentration. For this nutrient, the regression relationships gave satisfactory estimates for most compartments (mean error = 12–25%) and particularly for the stem.

? A procedure is proposed to estimate the nutrient exports associated with harvests of Pinus pinaster biomass.

     

Two-decadal trends in aboveground litterfall and net primary production in self-thinning Pinus banksiana stands in Wood Buffalo National Park,NWT, Canada

We estimated the aboveground net primary production (ANPP) in five self-thinning jack pine (Pinus banksiana) stands in Wood Buffalo National Park, NWT, Canada. The stands (11 to ca.175 years old) were selected to examine the relationship between stand density and tree size and its effect on carbon dynamics. Aboveground litterfall was collected from each stand from 1997 to 2012. Stand biomass was estimated by measuring tree size every 5 years and estimating the individual mass using allometric relationships. ANPP was then estimated by summing the litterfall mass, dead stem mass increment and stand biomass increment. We determined the proportional contribution of each organ to the total litter and the seasonal pattern of needle litterfall. There was a lower turnover rate of aboveground biomass in older stands than younger stands. The ANPP increased in the youngest stand (<30 years old) showed a decreasing trend in stands >50 years old. The maximum ANPP was estimated to be ca. 500?g m^?2 year^?1 in dry matter, which was found in 30–50 year-old stands.     

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.     

杨树新无性系冠层特性与生长关系研究

对 5年生黑杨无性系的冠层特性与材积生长之相关性进行了研究。材积生长与全树总叶面积TLA、树冠表面积TCA和冠型率CSR呈极显著正相关而与叶面积指数LAI、冠层密度CLD呈负相关。冠层内上、中层的叶面积对材积生长起了决定性作用 ,下层叶面积与材积生长关系不大。水平方向 ,冠层内、外部叶面积特性对材积生长贡献较大 ,而中部叶面积特性贡献小。阐明了杨树生长的理想冠层特性。     

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.     

麻竹枝叶生长对钩梢的响应

为了解冠层生物量积累、分配及枝叶大小对麻竹立竹受营林措施干扰(钩梢)后的响应,对不同竹龄全梢、钩梢麻竹地上构件生物量、生物量比、单叶特征、大小枝生物量分配比例及商品竹叶数量进行了调查.结果表明:麻竹立竹地上现存生物量分配格局为秆＞枝＞叶.叶生物量、叶/枝和叶/秆生物量比为2年生＞3年生＞1年生.随着竹龄的增加,枝、秆生物量、地上生物量和枝/秆生物量比总体上呈增加的趋势.2年生和3年生立竹枝、叶生物量分配比例显著高于1年生立竹,秆生物量分配比例显著小于1年生立竹.此外,随着竹龄的增加,立竹减少了对0 ～8 mm枝生物量的分配,增加了对8～16 mm、16 mm以上枝生物量的投入,以提高空间拓展能力,截获更多光资源.钩梢强烈影响了生物量分配格局,显著减小了麻竹立竹枝、叶、秆、地上生物量.钩梢后麻竹立竹增加了枝、叶生物量分配比例,减少了秆生物量的分配比例,同时提高了单叶叶面积和单叶干质量,增加了8～16 mm、16 mm以上枝生物量分配比例,减小0～8 mm枝生物量分配比例,以权衡枝叶的生长,提高立竹对环境的适合度.钩梢后立竹叶/枝、叶/秆、枝/秆生物量比升高,表明生物量分配更多地向叶和枝倾斜.钩梢麻竹商品竹叶数量较全梢麻竹增加29.68％,且发生部位明显降低,钩梢后冠层下部商品竹叶数量增加79.73％,中部商品竹叶数量增加25.81％,降低了采摘高度.在钩梢后的一个生长季内,钩梢影响了麻竹立竹资源利用策略,表现为枝与叶之间关系的变化,但随钩梢年限的增加其变化规律如何尚需进一步研究.     

Competition-density effect of tree organs in Acacia auriculiformis stands

The competition-density (C-D) effects for mean mass for tree, stem, branch and leaf were analyzed in Acacia auriculiformis stands. Mean tree mass-density and mean organ mass-density were well explained by the C-D equation of tree and the C-D equation of tree organ, respectively. An equation describing the relationship between mean leaf area u and density was formulated that fit the u-data well. The relationship between mean tree mass w and the ratio of each organ to mean tree mass (wo/ w) was examined. With increasing w, the stem mass ratio wS/w increased, whereas the branch mass ratio wB/w and the leaf mass ratio wL/w decreased. The yield difference between the lowest-density stand and the high-density stand became greater with stand growth. However, the yield of the mid-density stand was slightly lower than the yield of the high-density stand during the experimental period. To produce the most desirable combination of demanding individual-tree size and relative high stem yield, the mid-density is recommended as proper planting density for future management of A. auriculiformis stands.     

Both stem and crown mass affect tree resistance to uprooting

To examine the hypothesis that both stem and crown mass affect the resistance of a tree to uprooting and that tree resistance increases with increasing crown mass, we conducted tree-pulling experiments on three Picea glehnii plantations (stands A, B, and C: 27–32 years old) that differed in tree density and slenderness ratio. Allometries between crown and stem masses and between the critical uprooting moment and stem mass differed significantly among the three stands, with the crown mass and critical moment significantly larger in stand C than in stands A or B, despite the same stem mass. These results quantitatively verified our hypothesis. Allometries between crown and stem masses and between critical uprooting moment and stem mass were highly significant in each stand but were stand specific. Therefore, these allometries can be used to estimate tree resistance to uprooting in a given stand but not for data compiled from stands of various conditions and tree shapes. The allometry between critical moment and aboveground mass did not differ among the three Picea stands; thus, it is not stand specific and is generally appropriate to use for estimating tree resistance. To increase tree resistance to uprooting, we recommend light management for Picea glehnii plantations and probably other coniferous plantations as well.