Crown architecture of grafted Stone pine (Pinus pinea L.): shoot growth and bud differentiation

The singular umbrella-like crown shape of Stone pine can be interpreted as a consequence of primary shoot-growth patterns and posterior axis differentiation due to differential secondary growth and down-bending of branches. This paper centres on the first aspect, analysing the growth, branching and flowering behaviour of about 5,000 individual shoots on 27 grafted Stone pines. The data measurement on standing trees allowed to study correlations of topologic and geometric variables in the shoot and their ancestors. The only significant correlations were found with parameters of the mother shoot formed the previous year and with the number of cones born 3 years before by the respective ancestor. The fitted relationships between geometric and topologic shoot and branch variables are the first step of a structural model construction that can be completed with functional components like a radiation and a carbon allocation submodel, stressing the importance of the heavy Stone pine cones as carbon sinks, with a total annual allocation similar to stem wood. In conclusion, the Stone pine crown shape emerges as consequence of the lack of initial vigour differentiation between stem and main-branch apical meristems that favour the generalized sylleptic reiteration in the open-grown trees.     

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.     

秦岭南坡林窗大小对栓皮栎实生苗构型的影响

以秦岭南坡栓皮栎天然次生林间伐后形成的不同大小林窗实生苗为材料,采用典型抽样法调查林窗大小对栓皮栎实生苗枝系构型、叶片特征及其垂直分布的影响.结果表明: 林窗大小显著影响了栓皮栎实生苗的地径、树冠面积,林窗大小与地径呈显著正相关,与树冠面积呈显著负相关;林窗大小对实生苗苗高、冠长、树冠率均无显著影响;实生苗总体分枝率、逐步分枝率、枝径比均表现为大林窗>中林窗>小林窗>林下,不同林窗大小实生苗一级枝在垂直方向上主要分布于干的中、上部,直径较大的一级枝主要分布在中下部,一级枝倾角均随苗高的增高呈先增加后降低的趋势;随着林窗面积的增大,实生苗的叶长、叶宽和单叶面积逐渐〖JP2〗下降,平均单株叶数、相对高度上的总叶数增加,叶长宽比在不同大小林窗下均保持稳定,相对叶数均在干的中上部分布较多,叶面积指数与相对叶总数的变化一致;中林窗实生苗地径与大林窗无显著差异,但苗高较高,利于栓皮栎种群更新及培育优质干材.在栓皮栎林经营管理过程中,需确定合适的间伐强度增加中林窗数量,促进栓皮栎实生苗更新以及培育优质干材.     

短枝木麻黄种群苗期表型多样性评价

以来源于大洋洲原生种源区、亚洲原生种源区、亚洲引种次生区以及非洲引种次生区4个区域的20个种源短枝木麻黄种子和当年生幼苗为材料,通过种子千粒重以及幼苗苗高、地径、一级侧枝粗度、一级侧枝长度等7个性状对短枝木麻黄表型多样性进行了研究,以探讨种群苗期表型遗传差异,为短枝木麻黄早期遗传选择和遗传改良提供基本资料。结果表明:(1)短枝木麻黄种子千粒重在区域间和区域内种源间差异极显著,且千粒重具有显著的地理变异模式,随经度的增大而降低。(2)当年生幼苗苗高、地径在不同区域间及区域内种源间均存在极显著差异,其中泰国干东港种源幼苗生长最好(苗高76.6cm,地径4.64mm),而种源汤加的幼苗生长最差(苗高28.3cm,地径2.58mm)。(3)当年生幼苗一级侧枝粗度、一级侧枝长度、二级侧枝长度、每小枝节数和齿叶数在不同区域间及区域内种源间均存在极显著差异,其中齿叶数在区域间的变异系数最大(82.15%)。(4)通径分析表明,一级侧枝长度对苗高具有显著的正向影响作用,而一级侧枝粗度和二级侧枝长度对地径具有显著正向影响作用,它们可作为短枝木麻黄优良新品种筛选的参考因子。     

Maintenance mechanisms of the pipe model relationship and Leonardo da Vinci’s rule in the branching architecture of <Emphasis Type="Italic">Acer rufinerve</Emphasis> trees

The pipe model relationship (constancy of branch cross-sectional area/leaf area) and Leonardo da Vinci’s rule (equality of total cross-sectional area of the daughter branches and cross-sectional area of their mother branch) are empirical rules of tree branching. Effects of branch manipulation on the pipe model relationships were examined using five Acer rufinerve trees. Half the branches in each tree were untreated (control branches, CBs), and, for the others (manipulated branches, MBs), either light intensity or leaf area (both relating to photosynthetic source activity), or shoot elongation (source + sink activities), was reduced, and responses of the pipe model relationships were followed for 2 years. The pipe model relationship in MBs changed by suppression of source activity, but not by simultaneous suppression of source + sink activities. The manipulations also affected CBs in the year of manipulation and both branches in the next year. The branch diameter growth was most affected by light, followed by shoot elongation and leaf area, in that order. Because of the decussate phyllotaxis of A. rufinerve, one branching node can potentially have one main and two lateral branches. Analysis of 295 branching nodes from 13 untreated trees revealed that the da Vinci’s rule held in branching nodes having one shed branch but not in the nodes without branch shedding, indicating the necessity of natural shedding of branches for da Vinci’s rule to hold. These analyses highlight the importance of the source–sink balance and branch shedding in maintenance of these empirical rules. This article was contributed at the invitation of the Editorial Committee.     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

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。     

The occurrence of non-ring producing branches in Abies lasiocarpa

Lack of annual growth ring production at the base of branches in the lower portion of the crown is a commonly observed phenomenon. In this study, branches with missing rings were found on 47 of 49 Abies lasiocarpa trees sampled. The number of missing rings on the lowest live branch in the crown averaged 12, and ranged as high as 28. The lower one-third of the live crown tended to consist of branches without rings; these branches contained an average of 30% of the total crown leaf area. The ratio of foliage weight to total branch weight, in combination with position of the branch in the canopy, was an effective discriminator of non-ring producing branches. This suggests that both structural and environmental factors influence the cessation of ring production. The potential ecological implications of branches that fail to produce rings are discussed.     

不同栽培密度对辣木人工林分枝格局及生物量的影响

为了探讨干热河谷盆地辣木种植密度与分枝格局和生物量的关系及较为合理的种植密度,通过野外实地调查和数据分析对该地区不同栽培密度辣木人工林分枝格局及生物量进行研究,结果表明:辣木林分在株高、地径、冠幅以及分枝数量方面与栽培密度呈现显著负相关;不同密度下,根、茎、叶生物量及果荚数和果荚均长、直径和果荚均仔粒数之间都存在显著的差异性;栽培密度对辣木的分枝格局有较显著影响,主要表现在一级/二级分枝率、二级/三级分枝率以及三级分枝长度上有显著差异;辣木在不同方位上分枝格局有明显差异,主要体现在主、侧枝分枝长度和一级分枝角度上的差异显著;在本试验范围内,以叶用辣木为培育目标的较适宜的栽培密度为株行距0.9m×0.9m,以果用为栽培目标的较适宜的栽培密度为株行距1.5m×1.5m。在不同的栽培密度下,辣木人工林之间分枝格局及生物量的显著性差异,反映了辣木随环境的变化具有较高的形态可塑性和生态适应性。     

The hydraulic architecture of balsam fir (Abies balsamea)

Leaf-specific conductivities (LSCs – hydraulic conductivity per dry weight of supplied leaves). Huber values (transverse sapwood area per dry weight of supplied leaves), specific conductivity (hydraulic conductivity per transverse sapwood area) and tracheid diameters were measured throughout the trunk and crown of 20-year-old trees of Abies balsamca (L.) Mill. Measured specific conductivity was proportional to the radius to the fourth power of tracheids. LSCs, which indicate the relative water availability to different plant parts, are much higher in the trunk than in first order branches, and lowest in second order branches. The structural basis for this "hydraulic hierarchy" lies both in Huber values and in tracheid diameters. For similar diameter stem segments, there was no statistically significant difference for trunks versus branches in specific conductivity. However, in old parts of the tree, trunks are wider than supported branches and producer wider tracheids resulting in greater specific conductivities than in branches. In vigorous trees with strong apical control, Huber values were 12.0 times greater in the trunk than in similar diameter branch segments. In slow-growing trees with weak apical control, Huber values were 2.2 times greater in the trunk versus similar branch segments.     

孑遗树种钟萼木幼树生长特性探讨

研究了5 年生钟萼木幼树株高、地径连年生长、侧枝分生特性。结果表明,调查群体5 年生株高、地径平均值、最大值与最小值分别为2.0 m、3.4 m、0.5 m 和3.2 cm、9.5 cm、0.5 cm;群体幼树株高、地径生长分布遵循正态分布;幼树群体的株高、地径生长在早期表现显著的遗传分化。1～5 年生幼树的株高、地径年生长量表现为：株高在前3 年表现较低生长量,第4、5 年明显加快;地径在第1 年生长量占比较大。幼树株高小于2.0 m 的个体未发现分生侧枝,冠幅小且高生长较缓慢。     

Spatial variation in sapwood area to leaf area ratio and specific leaf area within a crown of silver birch

Spatial variation in sapwood area to leaf area ratio (Huber value, HV) and specific leaf area (SLA) was examined in branches of closed-canopy trees of silver birch (Betula pendula Roth). HV increased basipetally within a crown and decreased with increasing branch order, but exhibited no significant radial trend along a primary branch. HV was primarily determined by branch position in a crown and branch diameter at the sampling point, being independent of the size of the tree and branch. Greater HV in the lower-crown branches is considered a means to mitigate differences in hydraulic transport capacity between the branches located in different canopy layers. Beside branch position and sampling location on a branch, SLA depended significantly on several other variables characterising tree and branch size. SLA increased basipetally within a crown and along a primary branch, but exhibited no significant trend with branch orders. Because height caused leaf area (A_L) to diminish more rapidly than leaf dry weight, A_L primarily determined the vertical variation in SLA.     

Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

樟子松人工林一级枝条基径和枝长模型的研究

以东北林业大学帽儿山实验林场樟子松人工林为研究对象,采用枝解析的方法,于2002年、2003年测定了53株林木（年龄17~38 a,直径8.61~21.5 cm,树高7.48~18.24 m）的树冠变量,建立了基于总着枝深度（DINC）和树冠内一级枝条基径（BD）、枝长（BL）的预估模型。对于大小相同树木的一级枝条, 这些树冠变量随着DINC的增加而增大,而林木的胸径（DBH）、树高（HT）变量又很好地反映了不同大小树木的基径和枝长的变化。采用独立检验样本对构建的树冠内一级枝条基径和枝长模型进行了拟合统计量和精度检验,结果表明：模型预测效果良好,精度均达到95%以上。构建的一级枝条基径和枝长模型为进一步合理地描述樟子松人工林树冠的形状及其变化以及三维可视化经营提供依据。     

Growth pattern ofAbies mariesii saplings under conditions of open-growth and suppression

The growth pattern was analysed forAbies mariesii Mast. saplings in the subalpine climax forests of the northern Yatsugatake Mountains, Central Japan. The lateral branches were distinguished from the trunk not only with respect to the branching habits and needle arrangement but also with respect to the distribution of dry matter to needles, the slenderness of shoot-stem and the longevity of needles on them. Based on these differences, the contribution of the trunk and lateral branches to the matter production function of saplings is discussed. The lateral branches showed plastic changes in their growth characters in order to expand and maintain an assimilation system effectively, under conditions of suppression. In addition, the trunk growth was inhibited compared with the lateral branch growth and the apical dominance among lateral branches was weakened in the suppressed saplings. It was concluded that these growth habits resulted in the observed difference in the crown shape of saplings in the different habitats. Arguments are presented regarding the adaptive meanings of such a growth pattern for the shade tolerance and survival of the saplings and for the regeneration of the forest.     

山西濒危植物翅果油树植冠的构型分析

应用分形理论及方法,从分枝格局和冠幅扩展2个方面对山西翼城和乡宁的35株翅果油树(Elaeagnus mollis Diels)个体的植冠构型进行统计分析。结果表明,幼树和成树的总体分枝率和逐步分枝率有显著变化;一级枝的平均枝长和枝径有显著差异,但枝和叶的方位角及叶倾角差异不明显。不同发育阶段个体的冠幅变化较大,幼树冠幅的分形维数(2.0026)小于成树(2.2694)。翅果油树在不同生长发育阶段对生境变化有不同的构型策略。     

Ontogenic change in leaf shape and crown form of a tropical tree,Scaphium macropodum (Sterculiaceae) in Borneo

Crown architecture was analyzed forScaphium macropodum (Sterculiaceae), a common shade-tolerant emergent tree of a tropical rain forest in West Kalimantan, Indonesia. Saplings and poles shorter than 12 m in height had no branches, and gathered their leaves at the ends of the stem. The leaves changed from entire to palmately-parted with increasing tree size. The parted leaves increased the light penetration through the clustered foliage. The size of leaves including the blade and petiole ranged from 22 cm to 147 cm. Because the weight of petiole per blade increased with leaf size, the leaf could not be enlarged infinitely. Taller trees with lateral branches bore small (about 40 cm in length) entire leaves. The light intensity in the forest increased from the ground to about 12 m tall and was nearly constant from 12 m to 18 m. Crown architecture ofS. macropodum adapted to this light environment. The monoaxial trees lower than 12 m could thus increase the amount of light with vertical elongation, and the branched trees higher than 12 m could increase it by means of lateral extension of crown area.     

Cerambycid girdling and water stress modify mesquite architecture and reproduction

This study evaluated the effect of an outbreak of the cerambycid beetle, Oncideres rhodosticta, on branch growth and inflorescence production of the mesquite Prosopis glandulosa var. torreyana, and on larvae mortality in girdled branches at two sites (dry and wet) in the southern Chihuahuan Desert of Mexico. We compared stem growth responses to girdling in branches of similar sized trees from both sites over 42 months. The number of larvae per girdled branch was similar between sites, indicating similar ovipositing effort regardless of water stress. However, the proportion of dead larvae was significantly lower in trees at the dry site. On average, girdling reduced 96% of the stem length at both sites. At the end of the first year, 25% of the original stubs survived at the dry site, compared to 90% at the wet site. Girdling also broke apical dominance and allowed for the development of lateral buds from the surviving stubs, which produced many fewer branches and inflorescences at the dry site compared to the wet site, where a compact crown was developed. Water stress and girdling have a combined effect on mesquite architecture and reproduction, since more stubs died at the dry site while new branches from surviving stubs developed at the wet site, recovering the original lost biomass but changing the appearance of the tree.     

樟子松人工林分枝结构的分析

基于对6块樟子松(Pinus sylvestris var. mongolica)人工林固定标准地中的30株样木枝解析调查数据,通过分析不同林分、不同大小林木1级枝和2级枝的分枝概率、分枝格局和分枝角度,揭示了樟子松人工林树冠的分枝结构特点。研究结果表明：樟子松人工林1级枝和2级枝的平均分枝数量分别为3.84个和2.80个,两者分枝概率均呈正态分布;1级和2级枝条在光照条件好的几个区间（方位角46°～225°）分布较多,1级枝条的水平分布遵从均匀分布,而2级枝条则不遵从均匀分布;树冠上层枝条的分枝角度略小于树冠中、下层,上层平均分枝角度为45.6°,而中下层平均分枝角度都为49.4°。不同大小林木的1级枝分枝结构规律表明：Ⅰ级木和Ⅴ级木的每轮平均分枝数非常接近,分别为3.89和3.94个,比Ⅲ级木每轮分枝数大0.5个左右;1级枝水平分布在各区间内(45°间隔)相差在0.24%～2.81%之间,方差分析结果表明枝条水平分布与林木大小无关;不同大小林木的分枝角度有所差别,Ⅰ级木、Ⅲ级木和Ⅴ级木的平均分枝角度分别为48.5°、42.2°和50.7°。     

Structural crown properties of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in mixed versus pure stands revealed by terrestrial laser scanning

How tree morphology develops in mixed-species stands is essential for understanding and modelling mixed-stand dynamics. However, research so far focused on the morphological variation between tree species and neglected the variation within a species depending on intra- and interspecific competition. Our study, in contrast, addresses crown properties of nine mature Norway spruces (Picea abies [L.] Karst.) of a pure stand and compares them with ten spruces growing in mixture with European beech (Fagus sylvatica [L.]). The same was done with 11 pure stand beeches and 12 beeches growing in mixture with spruce. Through application of a terrestrial laser scanner and a new skeletonization approach, we deal with both species’-specific morphological traits such as branch angle, branch length, branch bending, crown volume and space occupation of branches within the crown, some of which were hardly accessible so far. Special attention is paid to distinct differences between trees growing in mixed and pure stands: for spruce, our study reveals significantly longer branches and greater crown volumes in the mixed stand when compared to the pure stand. In case of European beech, individuals growing in mixture show flatter branch angles, more distinct ramification, greater crown volumes and a lower share of a single branch’s space occupation in the total crown volume. The results show that the presented methods yield detailed information on the morphological traits analyzed in this study and that interspecific competition on its own may have a significant impact on crown structures. Implications for production ecology and stand dynamics of mixed-species forests are discussed.