Development of biomass proportions in Norway spruce (Picea abies [L.] Karst.)

This study tests the hypotheses that (1) the above-ground structure of Norway spruce (Picea abies [L] Karst.) is derivable from the functional balance theory, and that (2) crown ratio is a key source of structural variation in trees of different age and social position. Twenty-nine trees were measured in three stands (young, middle-aged, and mature), with three thinning treatments (unthinned, normal, and intensive) in the two older stands. There was a strong linear relationship between the total cross-sectional area of branches and that of stem at crown base. Foliage mass was linearly related with stem basal area at crown base. Also an allometric relationship was found between foliage mass and crown length. The mean length (weighted by basal area) of branches obeyed an exponential function of crown length. The parameters of most of these relationships were independent of slenderness (tree height/breast height diameter) and tree age However, total branch cross-sectional area per stem cross-sectional area in the young trees was greater than in the older trees. The young trees also had slightly shorter branches than predicted by the mean branch length equation. This was probably caused by branch senescence which had not yet started in the young stand. The older trees had a relatively long lower crown segment which was growing slowly and senescing. It was proposed that a segmented crown structure is characteristic of shade tolerant tree species, and that the structural model could be further developed by making the two segments explicit.     

Effects of age and site quality on the distribution of biomass in Scots pine (Pinus sylvestris L.)

The distribution of the above-ground and below-ground biomass of Scots pine in southern Finland were investigated in trees of different ages (18–212 years) from two types of growth site. Secondly, some structural regularities were tested for their independence of age and growth site. Trees were sampled from dominant trees which could be expected to have a comparable position in stands of all ages. All stands were on sorted sediments. The biomass of the sample trees (18 trees) was divided into needles, branch sapwood and heartwood, stem sapwood and heartwood, stem bark, stump, large roots (diameter >20 cm), coarse roots (five classes) and fine roots. The amount of sapwood and heartwood was also estimated from the below-ground compartments. Trees on both types of growth site followed the same pattern of development of the relative shares of biomass compartments, although the growth rates were faster on the more fertile site. The relative amount of sapwood peaked after canopy closure, coinciding with the start of considerable heartwood accumulation. The relative amount of needles and fine roots decreased with age. The same was true of branches but to a lesser degree. The relative share of the below-ground section was independent of tree age. Foliage biomass and sapwood cross-sectional area were linearly correlated, but there were differences between the growth sites. Needle biomass was linearly correlated with crown surface area. The fine root to foliage biomass ratio showed an increasing trend with tree age.     

Factors influencing choice of balsam fir twigs from thinned and unthinned stands by moose

Summary Moose were observed to browse preferentially on balsam fir (Abies balsamea) trees in stands where the stem density had been mechanically reduced to about 2000 stems/ha from over 30,000 stems/ha. Twigs from trees in thinned and unthinned stands were analyzed to test the hypotheses that moose were choosing thinned stands to maximize intake of a nutrient, or avoiding plant secondary compounds deliterious to digestion. Analyses included: twig length, weight, diameter, acid detergent fiber, neutral detergent fiber, dry matter digestibility, macroelements, crude fat (resins), crude protein, ash, and volatile secondary metabolites. Significantly higher concentrations of crude protein, P, Ca, Na, and crude fats occurred in trees from thinned compared to unthinned stands. Twigs from thinned stands were more digestible, longer, heavier, and had a greater diameter than those from unthinned stands. Several secondary metabolites were found in highest concentration in thinned stands. We suggest that moose chose thinned stands over unthinned stands for feeding because of high protein levels and large twigs. Trees in unthinned stands had protein levels below reported maintenance levels required by moose. Secondary metabolite concentrations were opposite to the direction predicted and the data do not support plant defence hypotheses for the chemicals analyzed.     

Diversity and succession of epiphytic macrolichen communities in low‐elevation managed conifer forests in Western Oregon

Abstract. We examined epiphytic macrolichen communities in Pseudotsuga menziesii (Douglas‐fir) forests across the western Oregon landscape for relationships to environmental gradients, stand age and structure, and commercial thinning. We used a retrospective, blocked design through the Coast and the western Cascade ranges of Oregon. Each of our 17 blocks consisted of a young, unthinned stand (age 50–110 yr); an adjacent, thinned stand of equivalent age; and an old‐growth stand (age > 200 yr). We found 110 epiphytic macrolichen taxa in the stands. Forage‐providing alectorioid lichens and the nitrogen‐fixing cyanolichen Lobaria oregana associated strongly with old‐growth stands and remnant old trees in younger stands (unthinned + thinned). Relative to unthinned stands, thinned stands had a slightly higher abundance of alectorioid lichens and a greater presence of Hypogymnia imshaugii. However, thinned stands hosted a lower landscape‐level (γ) diversity, lacking many species that occurred infrequently in the unthinned stands. Patterns in the lichen community composition correlated strongly with climatic gradients; the greatest variation in composition was between the Coast and Cascade ranges. The difference in communities between mountain ranges was greatest among stands 70–110 yr old, suggesting a difference in lichen successional dynamics between the ranges.     

Relationship between an exotic phloem feeder and balsam fir (Abies balsamea L. (Mill.)) foliar chemistry: implications for two native defoliators

The balsam woolly adelgid (Adelges piceae) is a gout-inducing hemipteran native to the silver fir forests of Europe. Introduced to eastern North America approximately 100 years ago, it is now found in most balsam fir forests in Atlantic Canada. When A. piceae feed, they trigger a reaction in the host branch that alters both xylem and phloem morphology. We conducted a field survey to examine the relationship between A. piceae gout density and balsam fir foliar chemistry and shoot growth in naturally unthinned and precommercially thinned stands. A. piceae gout density negatively affected branch growth and was related to changes in the chemistry of older, but not current-year foliage. Older foliage experienced decreases in camphene and bornyl acetate, while foliar concentrations of camphene, myrcene, phenolics, potassium and water differed between thinned and unthinned stands. Foliar chemistry was also influenced by interactions between thinning and A. piceae gout density in old foliage. This study suggests that changes in balsam fir associated with A. piceae gout density may force native defoliators that feed in highly gouted trees to adapt to diets of different chemical compositions and that thinning may alter these interactions.     

Crown profile of foliage area characterized with the Weibull distribution in a hinoki (Chamaecyparis obtusa) stand

Summary Thirten sample trees of various sizes in a 29-year-old hinoki [Chamaecyparis obtusa (Sieb, et Zucc.) Endl.] plantation were felled and subjected to the stratified clip technique. Crown profile of foliage area fitted well with the Weibull distribution. The crown profile tended to be more skewed toward the top of crowns in smaller trees than in larger trees. This tendency was reflected in the value of the shape parameter of the Weibull distribution. The shape parameter ranged from 1.73 to 3.23 and gradually increased up to an asymptotic value with an increase of stem diameter at breast height. The scale parameter of the distribution ranged from 1.0 to 4.2 and tended to increase in proportion to stem diameter at breast height. Foliage area of a tree correlated well with stem diameter at breast height through an ordinary allometric equation. Tree height could be approximated fairly well by a generalized allometric equation as a function of stem diameter at breast height. On the basis of the census of stem diameter at breast height, canopy profile could be constructed synthesizing crown profiles of foliage area for individual trees in the stand. Leaf area index was estimated to be 6.6 ha ha^–1.     

Development of branch,crown, and vertical distribution leaf area models for contrasting hardwood species in Maine,USA

Key message

Branch, crown vertical leaf area distribution models were developed for naturally regenerated hardwood species and planted hybrid poplar clones. Species-specific differences were found at all levels of investigation.

Abstract

Coexistence in mixed-species stands is strongly influenced by species differences in leaf area production and distribution. The majority of leaf area models in the literature are focused on conifer species, which have substantially different crown forms than hardwood species. Therefore, the goal of this investigation was to develop branch, crown, and vertical leaf area distribution models for various hardwood species that accounted for their greater crown complexity. A nonlinear model including branch diameter, branch tip height, and height to the start of the foliage was the best fit for branch leaf area. Branch leaf area ranged from 0.05 to 0.37 m² for Populus grandidentata and Betula populifolia for an averaged sized branch, respectively. The best fit model for crown leaf area was a nonlinear form accounting for stem diameter and crown length. Crown leaf area ranged from 3.26 to 9.85 m² for Populus tremuloides and Betula populifolia for an averaged sized tree, respectively. Vertical leaf area distribution was best fit by a right-truncated Weibull distribution and showed a peak in the middle third of the crown for most of species. In addition, leaf area production varied among four hybrid poplar clones in plantations, suggesting a strong genetic control over crown form. Overall, leaf area varied among species at all levels of investigation, suggesting that coexistence of hardwood saplings in this investigation was strongly influenced both by inherent species-specific leaf area production and vertical distribution.     

Competition-dependent modelling of foliage biomass in forest stands

Currently, foliage biomass is estimated based on stem diameter or basal area. However, it is questionable whether the relations between foliage and stem observed from plantations of a single tree species can be applied to stands of different structure or species composition. In this paper, a procedure is presented to simulate foliage and branch biomass of tree crowns relative to crown size and light competition. Crowns are divided into layers and segments and each segment is divided into a foliated and an unfoliated fraction. Depending on the competitive status of the segment, leaf area density, specific leaf area and foliated branch fraction are determined. Based on this information, foliage biomass is calculated. The procedure requires a crown shape function and a measure to characterise competition for light and space of each individual segment within the canopy. Simple solutions are suggested for both requirements to enable an application with data that can be measured non-destructively in the field; these were stem position, tree height, crown base height, crown radii and some general crown shape information. The model was parameterised from single trees of Norway spruce and European beech and partly evaluated with independent data close to the investigation plot. Evaluations showed that the model can attribute the ecology of the different crown forms. Modelled foliage distribution for beech and spruce as well as total needle biomass of spruce agreed well with measurements but foliage biomass of beech was underestimated. The results are discussed in the context of a general model application in structured forests.     

樟子松人工林树冠结构的分形分析

基于樟子松人工林7块固定标准地中的31株解析木的树冠体积和叶量,以幂函数关系(F=Av^(D/3))建立了预估树冠表面积的分形维数。同时根据生物量实测数据,建立预估叶量的生物模型Lw=0.180397D^3045903H^-1.67348。基于枝解析、树干解析数据,动态地预估了一年、二年、三年前的树冠体积,并结合树冠体积、叶量的这种幂函数关系可以动态地预估一年、二年、三年前树冠表面积的分形维数,从而反映出树冠结构的动态变化规律。为了了解不同分级样木的分维数变化情况,利用2003年调查的4块生物量标准地数据,根据单株树木各个枝条占据的空间体积与该枝条的带叶枝干重的关系,计算了各标准地不同分级样木树冠的分维数。为探讨单株样木树冠的分维数的计算提供了一种可行方法。树冠的分维数作为表征树冠的动态生长变化是一有用和可靠的指标。     

Modelling the distribution of diameter growth along the stem in Scots pine

This paper presents an empirical model for the distribution of diameter growth along the stem in Scots pine (Pinus sylvestris L.) and for the consequent stem form over time. First, the distribution of annual mass growth in the stem is determined as a function of the total annual growth in stem mass, current stem mass and the distribution of the latter along the stem. Second, the distribution of diameter growth is obtained by converting the fraction of annual growth in the stem mass at a given height in the stem into the thickness of the annual ring at the same height. Application of the model to Scots pine data sets including both young and mature trees not used in parameter estimation showed that the model was capable of reconstructing the distribution of diameter growth from the stem butt to the apex and from the pith to the stem surface at any height in the stem in both young and mature trees. The resulting empirical model was also linked to a physiological, process-based model in order to study its performance in a simulated stand. Simulations representing trees grown in unthinned and thinned Scots pine stands with trees of different status (from dominant to suppressed) showed that the response in tree growth to thinning in terms of the distribution of diameter growth along the stem was quite realistic relative to measured data.     

Specific leaf area of European Larch (<Emphasis Type="Italic">Larix decidua</Emphasis> M<Emphasis Type="SmallCaps">ill</Emphasis>.)

Key message

The specific leaf area of European larch depends on branch height and canopy depth, indicating that both, the effect of hydraulic limitations and low water potentials in greater branch heights, and light availability affect specific leaf area.

Abstract

Specific leaf area (SLA) is defined as the ratio between projected leaf area and needle dry mass. It often serves as parameter in ecosystem modelling as well as indicator for potential growth rate. We explore the SLA of European larch (Larix decidua) and the most important factors which have an influence on it. Data were collected from eight stands in Styria, Austria. The stands varied in age, elevation and species mixture. Four stands were pure larch stands with only minor proportions of Norway spruce (Picea abies), whereas the other four were mixed stands of larch and spruce. In each stand 15 representative sample trees were felled. The crown of each sample tree was divided into three sections of equal length and in each section a random sample of needles was taken for determining projected leaf area and dry mass of 50 needles. The mean SLA of larch was established to be 117 cm² g^?1 with a standard deviation of ±27.9 cm² g^?1. SLA varies within the crown, but neither between different mixtures nor years of observation nor social position of the trees. A mixed-effects model, with the plots as random effect, revealed that SLA of larch decreased with increasing branch height (p = 0.0012) and increased with increasing canopy depth (p = 0.029). We conclude that both the hydraulic limitations due to low water potentials in greater branch heights and light availability affect specific leaf area.

     

Seasonal prevalence of arthropods after line thinning of overstocked Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantations in central Japan

We investigated the seasonal changes in the abundance of coleopteran and hymenopteran functional groups to evaluate the ecological suitability of line thinning in overstocked Japanese cedar plantations in central Japan. In line thinning, 3- to 5-m-wide sections of the stand are harvested parallel to the mountain slope and perpendicular to the forest roads. The thinning intensity varies from 25 to 35% of the total number of trees. We compared the seasonal abundance between the thinned stand and the unthinned (control) stand in two plantation forests: a low-elevation Sugi site (4 years since thinning) and a high-elevation Kuchiotani site (6 years since thinning). Most coleopteran and hymenopteran functional groups were consistently more abundant in the thinned stand than in the unthinned stand. The differences in arthropod abundance between the thinned and unthinned stands were mainly caused by marked differences in the abundance of seasonally strongly changing functional groups during their peak seasons. The predators and parasitoids of insects as well as the plant and pollen feeders (plant-dependent functional groups) responded to seasonal changes in the vegetation (food and microhabitats) and microclimate. The ants were less affected by line thinning. Our results indicate that line thinning is an ecologically suitable silvicultural treatment for the improvement of biodiversity conditions in the overstocked plantation forests.     

Size structure of current-year shoots in mature crowns

Characteristics of current-year shoot populations were examined for three mature trees of each of three deciduous broad-leaved species. For first-order branches (branches emerging from the vertical trunk) of the trees examined, lengths or diameters of all current-year shoots were measured. Total leaf mass and total current-year stem mass of first-order branches were estimated using an allometric relationship between leaf or stem mass and length or diameter of current-year stems. For each tree, the number of current-year shoots on a first-order branch was proportional to the basal stem cross-sectional area of the branch. On the other hand, first-order branches had shoot populations with size structures similar to each other. As a result, the leaf mass of a first-order branch was proportional to the basal stem cross-sectional area of the branch, being compatible with the pipe-model relationship. All current-year shoot populations had positively skewed size structures. Because small shoots have a larger ratio of leaf mass to stem mass than large shoots, first-order branches had an extremely large ratio of leaf mass to current-year stem mass. This biased mass allocation will reduce costs for current stem production, respiration and future radial growth, and is beneficial to mature trees with a huge accumulation of non- photosynthetic organs. The allometric relationships between leaf mass and basal stem diameter and that between leaf mass and current-year stem mass of first-order branches were each similar across the trees examined. Characteristics of shoot populations tended to offset inter-species diversity of shoot allometry so that branch allometry shows inter-species convergence.     

Understanding changes in black (Picea mariana) and white spruce (Picea glauca) foliage biomass and leaf area characteristics

Key message

Growth conditions related to inter-tree competition greatly influence black and white spruce foliage biomass and projected leaf area characteristics.

Abstract

Foliage characteristics such as biomass and area are important among other reasons because they can be related to tree growth. Despite their economic and ecologic importance, equations to characterize foliage biomass and projected area of black (Picea mariana (Miller) BSP) and white (Picea glauca (Moench) Voss) spruces are sparse. Total foliage biomass and projected leaf area, foliage biomass and leaf area density, and relative vertical distribution of black and white spruces foliage biomass and leaf area were modelled with linear and nonlinear mixed effect models. A total of 65 white spruces and 57 black spruces were destructively sampled at four different locations in Alberta, Québec, and Ontario, Canada. Our results show that for each species, total tree foliage biomass and projected leaf area is proportional to stem diameter, total height, and crown length. The addition of crown length in the equations improved the precision of the predictions of total foliage biomass for both species and diminishes greatly the site level random effect. An increase in DBH for black spruce and in the DBH to total height ratio for white spruce skewed the relative vertical foliage biomass distribution toward the base of the living crown. According to our results, growth conditions or tree development stage influence both foliage biomass and leaf area characteristics of black and white spruces. Our results emphasize the importance of inter-tree competition on foliage biomass characteristics.     

不同密度红桦幼苗苗冠结构与竞争对CO2浓度升高的响应

研究了两个种植密度下,红桦（Betula albosinensis）苗冠结构特征对CO2浓度的响应,在此基础上探讨了CO2浓度升高对植物竞争压力的影响。结果表明,冠幅、冠高、苗冠表面积和苗冠体积均受CO2浓度升高的影响而增加,但是受密度增加的影响而降低。CO2浓度升高对苗冠的促进效应在低密度条件下大于高密度处理,高密度条件下苗冠基本特征部分地受到CO2浓度升高的促进作用;升高种植密度的效应则在高CO2浓度条件下大于现行CO2浓度处理。高CO2浓度和高密度条件下,LDcpa（单位苗冠投影面积叶片数）、LDcv（单位苗冠体积叶片数）和苗冠底部枝条的枝角均低于相应的现行CO2浓度处理和低密度处理,这主要是由于冠幅和冠高的快速生长所造成的。升高CO2浓度对枝条长度的影响与枝条在主茎上所处位置有关。总之,升高CO2浓度有利于降低增加种植密度对苗冠所带来的负效应,而增加种植密度降低了升高CO2浓度的正效应。LDcpa和LDcv的降低表明,红桦在升高CO2浓度和种植密度的条件下,会作出积极的响应,从而缓解由于生长的增加所带来的竞争压力的增加。     

Distribution of vessel size,vessel density and xylem conducting efficiency within a crown of silver birch (<Emphasis Type="Italic">Betula pendula</Emphasis>)

Spatial patterns in vessel diameter, vessel density and xylem conducting efficiency within a crown were examined in closed-canopy trees of silver birch (Betula pendula). The variation in anatomical and hydraulic characteristics of branches was considered from three perspectives: vertically within a crown (lower, middle and upper crown), radially along main branches (proximal, middle and distal part), and with respect to branch orders (first-, second- and third-order branches). Hydraulically weighted mean diameter of vessels (D _h) and theoretical specific conductivity of the xylem (k _t) exhibited no vertical trend within the tree crown, whereas leaf-specific conductivity of the xylem (LSC_t) decreased acropetally. Variation in LSC_t was governed by sapwood area to leaf area ratio (Huber value) rather than by changes in xylem anatomy. The acropetal increase in soil-to-leaf conductance (G _T) within the birch canopy is attributable to longer path length within the lower-crown branches and higher hydraulic resistance of the shade leaves. D _h, k _t and LSC_t decreased, while vessel density (VD) and relative area of vessel lumina (VA) increased distally along main branches. A strong negative relationship between vessel diameter and VD implies a trade-off between hydraulic efficiency and mechanical stability of xylem. D _h and VD combined explained 85.4% of the total variation of k _t in the regression model applied to the whole data set. Xylem in fast-growing branches (primary branches) had greater area of vessel lumina per unit cross-sectional area of sapwood, resulting in a positive relationship between branch radial growth rate and k _t. D _h, k _t and LSC_t decreased, whereas VD increased with increasing branch order. This pattern promotes the hydraulic dominance of primary branches over the secondary branches and their dominance over tertiary branches. In this way crown architecture contributes to preferential water flow along the main axes, potentially providing better water supply for the branch apical bud and foliage located in the outer, better-insolated part of the crown.     

Sapwood hydraulic conductivity and leaf area – sapwood area relationships following thinning of a Eucalyptus nitens plantation

This study tested the hypothesis that the relationship between leaf area (projected, or one‐sided) and sapwood area in Eucalyptus nitens (Deane and Maiden) Maiden is affected by thinning treatment. However, no difference in the relationship between leaf area and sapwood area was found 8 years after thinning. This result suggests that a single regression equation can be used to predict the leaf area of trees in thinned and unthinned stands. The relationship was non‐linear, implying a causal relationship between growth rate and the ratio of leaf area to sapwood area (A_l : A_s). Sapwood hydraulic conductivity increased by approximately 100% from breast height to crown base, whereas sapwood area decreased by 19%. This implies that the efficiency of water transport through the sapwood increased by 60% along this length. This conclusion is supported by the A_l : A_s relationships which showed that the sapwood area at crown base supported, on average, close to 60% more leaf area than a similar amount of sapwood area at breast height. That large trees in this study had greater hydraulic conductivity and higher A_l : A_s lends support to the argument that resource capture, and hence growth rate, influence sapwood hydraulic conductivity.     

北京地区侧柏人工林密度效应

密度是影响森林尤其是人工林生长的重要因素,林冠层是森林生态系统与其他系统进行能量和物质交换的重要场所,树木及树冠生长对林分密度的响应关系可以看作是生物对环境变化产生的适应性现象。林分密度效应是生态学和森林培育学的重要研究内容之一。以23块8种不同密度梯度的北京山区侧柏人工幼龄林林分为研究对象分析其树木生长及树冠生长对密度的响应关系,其中树冠指标使用了参照了美国林务局(USDA)的树冠调查指标。研究结果表明:(1)林分平均胸径、平均树高和平均冠幅生长均随密度增大而减小,林分密度大于3000株/hm2时各指标减小的趋势变缓,使用异速生长模型可以很好地拟合这种变化关系;(2)随密度增加,树冠水平方向和垂直方向生长均到显著地抑制作用,树冠外形表现出由饱满冠型向狭长冠型变化的适应性现象;(3)使用树冠二维、三维指标与密度进行相关性分析可知树冠长度、树冠率等指标与林分密度呈负相关关系,树冠圆满度及树冠生产效率与密度表现出极显著正相关关系;(4)采用枝解析的方法研究了树枝长度、材积的平均生长量、连年生长量与密度的关系,结果表明幼龄期各生长量差异不大;(5)在建立冠幅模型时考虑了自变量间的多重共线性问题,所建的胸径单自变量二次方模型能够很好地预测侧柏人工幼龄林冠幅生长过程,模型相关系数R2为0.961。     

Implications of varying pipe model relationships on Scots Pine growth in different climates

1. The process-based model SIMFORG , based on the pipe theory, was parameterized for Scots Pine at six locations along a north–south gradient in Europe. The ratio of foliage mass to stem cross-sectional area was changed as a function of potential evapotranspiration as proposed by Berninger et al. (1995).
2. Allocation to the stem differed between the locations and affected consequently the stemwood production. Variation in the net primary production and differences in the pipe model parameters were responsible for these differences. There was good agreement between measured and simulated data.
3. Increase in primary production, as predicted by climate-change senarios, increased allocation to the stem. However, the results were sensitive to changes in the foliage mass to stem sapwood cross-sectional area ratio. The changes in allocation were higher in the north than in the south.     

Effects of individual size,local competition and canopy closure on the stem volume growth in a monoclonal Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantation

We studied how the dominant factor affecting stem volume growth changes during stand development in a monoclonal stand of Cryptomeria japonica D. Don. Stem analysis was used to compare growth history of trees in an unthinned plot (closed canopy) and a thinned plot (open canopy). In the unthinned plot, the dominant factor affecting stem volume growth was basal area (BA) before canopy closure, whereas neighborhood competition index (CI) was the dominant factor after canopy closure. In contrast, the dominant factor affecting stem volume in the thinned plot was BA throughout stand development. Spearman’s rank correlation coefficient between BA and CI continued to increase after canopy closure and size rank among individuals became increasingly fixed. Our results indicated that stem volume growth shifts from size-dependent to competition-dependent growth at canopy closure. The apparent correlation between tree size and growth rate observed in many previous studies may be the result of competition-mediated positive feedback between size and growth.