The anatomy and grain pattern in forks of hazel (Corylus avellana L.) and other tree species

Key message

This paper provides a new anatomical model for how branches are joined together at forks in trees, using evidence from MicroXCT scanning, ESEM output and visual observation.

Abstract

Wood grain arrangements at the forks and other junctions within a tree must be arranged to mechanically join together the two or more branches, yet not adversely restrict sap flow. The grain orientation at junctions therefore represents a trade-off in xylem performance between the functions of efficient sap conductance and the provision of adequate load-bearing capacity. Initial observations of wood grain orientation were made on the surfaces of several dozen debarked and fractured bifurcations of a wide range of tree species, both by eye and using a scanning electron microscope. Subsequently, small volumes of wood were sampled from two locations within the junctions of hazel, at the junction apex and on the outer section of join. Wood was imaged in 3D using high-resolution X-ray tomography, and the scanned volumes were analysed for their wood grain patterns. It was found that the wood at the junctions of hazel forks contained only 37 % of the number of vessels contained in wood within the adjacent stem. The vessel elements formed at the junctions were only 32.5 % the length of those in the stem, had a mean diameter only 50.5 % of the stem vessels and consequently only 26.3 % of their lumen volume. The passage of the vessels through the fork wood deviated from a straight line (Euclidean) distance by more than 14 times more than the stem wood vessels did. The interweaving of vessels in the fork wood was over 22 times greater than in the stem wood. A survey of rays showed them to be 58 % more abundant in fork wood but only 62 % of the height of rays in the stem wood. These results suggest that where two branches of similar diameter join to form a tree fork, an interlocking wood grain pattern is formed at the apex of the bifurcation, which provides higher tensile strength to the junction. Breaking of the hazel fork requires wood fibres to be stretched axially and broken across, which requires greater stress than breaking normal wood radially or tangentially.     

A mathematical model of auxin-mediated radial growth in trees

A mathematical model is presented to describe the coupling between the concentration of indole-3-acetic acid (IAA) in the cambial region of a tree branch and the radial expansion of the branch during active growth. The main features of the model are (1) the branch cambium is treated as an approximately cylindrical surface of negligible thickness, (2) the rate of radial growth is proportional to the mass of IAA per unit area on the cambial surface, and (3) IAA is transported basipetally through the cambium at a constant speed. We neglect the role of elastic strains in the determination of branch shape, and the effects of IAA synthesis and metabolization in the cambium, so the model is not quantitatively accurate. However, the model does reproduce several important qualitative features of tree growth including the approximate area-preserving property of tree branch junctions and the ability of a branch to maintain its shape despite perturbations due to injury.     

Autoinhibition of indoleacetic acid transport in the shoots of two-branched pea (Pisum sativum) plants and its relationship to correlative dominance

Two-branched pea plants ( Pisum sativum L. cv. Lisa ZS) with different dominance degrees, obtained by removing the epicotyl shortly after germination, were used to study the interaction between the polar transport of indoleacetic acid (IAA) in both branches of the plants and its relationship to correlative dominance. The dominant shoot had higher transport capacity for 3H-IAA, exported more IAA out of its apex and possessed more endogenous IAA in apex and the first internode than the dominated one. Decapitation of the dominant shoot resulted in a rapid resumption of growth in the dominated shoot, accompanied by a considerable increase in its capacity to export endogenous IAA and to transport 3H-IAA. Parallel experiments with intact two-branched plants and Y-formed explants showed that the 3H-IAA transport on one side was inhibited by the other branch apex or by pre-application of 12C-IAA to the cut stump of the decapitated side. The higher the concentration of 12C-IAA applied to the cut stump of one side of the Y-form explant was used, the stronger the 3H-IAA transport was inhibited and the more the transported IAA was conjugated above the junction on the other side. The results of these experiments support the autoinhibition hypothesis at junctions. The relationship between elongation growth and IAA export/transport in the two-branch pea plants is considered.     

Patterns of auxin distribution during gravitational induction of reaction wood in poplar and pine

Gravistimulation of tree stems affects wood development by unilaterally inducing wood with modified properties, called reaction wood. Commonly, it also stimulates cambial growth on the reaction wood side. Numerous experiments involving applications of indole-3-acetic acid (IAA) or IAA-transport inhibitors have suggested that reaction wood is induced by a redistribution of IAA around the stem. However, in planta proof for this model is lacking. Therefore, we have mapped endogenous IAA distribution across the cambial region tissues in both aspen (Populus tremula, denoted poplar) and Scots pine (Pinus sylvestris) trees forming reaction wood, using tangential cryosectioning combined with sensitive gas chromatography-mass spectrometry analysis. Moreover, we have documented the kinetics of IAA during reaction wood induction in these species. Our analysis of endogenous IAA demonstrates that reaction wood is formed without any obvious alterations in IAA balance. This is in contrast to gravitropic responses in roots and shoots where a redistribution of IAA has been documented. It is also of interest that cambial growth on the tension wood side was stimulated without an increase in IAA. Taken together, our results suggest a role for signals other than IAA in the reaction wood response, or that the gravitational stimulus interacts with the IAA signal transduction pathway.     

Natural and experimentally altered hydraulic architecture of branch junctions in Acer saccharum Marsh. and Quercus velutina Lam. trees

 The functional xylem anatomy and the hydraulic conductivity of intact and treated branch junctions of the diffuse-porous sugar maple (Acer saccharum Marsh.) were compared to those of the ring-porous black oak (Quercus velutina Lam.). Maple shoots possessed greater growth intensity than those of oak. The extensive growth of the maple trees resulted in about a two-fold increase in xylem production in the maple branches. Branches were altered by removing a patch of bark from the base of a branch (near a junction) leaving a bridge of bark on the upper or lower side of the branch. The experimentally treated branch junctions revealed that, in oak, most (up to 92%) of the water flows in the lower side of a branch, where most of the large vessels occurred. In maple, most of the conductive tissue was observed to form in the upper side of the branches, which was equally or more conductive than the lower side. A treatment of longitudinal, parallel scratches in the bark-bridge, which reduced earlywood vessel width, substantially decreased conductivity (to only 15%) in oak, but had no effect on conductivity in maple. In maple, such wounding stimulated more wood formation and increased conductivity. In both trees, a narrow bridge at the junction induced more wood formation and higher conductivity in the branch. The mechanisms controlling wood formation and water flow in branch junctions of ring- and diffuse-porous trees are discussed. Received: 14 February 1996 / Accepted: 27 May 1996     

Whole-tree water transport scales with sapwood capacitance in tropical forest canopy trees

The present study examines the manner in which several whole‐tree water transport properties scale with species‐specific variation in sapwood water storage capacity. The hypothesis that constraints on relationships between sapwood capacitance and other water relations characteristics lead to predictable scaling relationships between intrinsic capacitance and whole‐tree behaviour was investigated. Samples of sapwood from four tropical forest canopy tree species selected to represent a range of wood density, tree size and architecture, and taxonomic diversity were used to generate moisture release curves in thermocouple psychrometer chambers, from which species‐specific values of sapwood capacitance were calculated. Sapwood capacitance was then used to scale several whole‐tree water transport properties determined from measurements of upper branch and basal sap flow, branch water potential, and axial and radial movement of deuterated water (D₂O) injected into the base of the trunk as a tracer. Sapwood capacitance ranged from 83 to 416 kg m^?3 MPa^?1 among the four species studied and was strongly correlated with minimum branch water potential, soil‐to‐branch hydraulic conductance, daily utilization of stored water, and axial and radial movement of D₂O. The species‐independent scaling of several whole‐tree water transport properties with sapwood capacitance indicated that substantial convergence in plant function at multiple levels of biological organization was revealed by a simple variable related to a biophysical property of water transport tissue.     

Biomechanics of a branch – stem junction in softwood

Direct measurement of strain field in a mechanically loaded Norway spruce branch–stem junction was performed by means of electronic speckle pattern analysis. Results were compared with strain distribution in a polyester cast of identical shape as the branch–stem junction, and a simplified polyester model consisting of two half-cylinders. Compared to polyester models, the branch–stem junction was characterised by a very homogeneous distribution of strain, which can be interpreted as a homogeneous distribution of stress in terms of fraction of material strength. This optimised transfer of mechanical load from the branch to the stem is achieved by a combination of naturally optimised shape with, additionally, optimised mechanical wood properties in the junction area.     

Wood Grain Pattern Formation: A Brief Review

In trees, new wood develops from a layer of stem cells called the vascular cambium. A subpopulation of cambial cells—the fusiform initials—are elongated and capable of coordinated reorientation in response to internal and external stimuli. Changes in the orientation of fusiform initials in turn leads to changes in the grain pattern of developing wood. This article reviews the phenomenon of cambial orientation, with an emphasis on a recent computer model that takes the plant hormone auxin as the orienting signal. New model results are presented that demonstrate the surprisingly complex grain patterns that can emerge from simple initial conditions, in qualitative agreement with similar patterns found in wood. Lastly, an alternative theory of wood grain pattern that takes mechanical stress as the orienting signal is critically evaluated.     

基于地基激光雷达的落叶松人工林枝条因子提取和建模

地基激光雷达(TLS)可以实现从森林中无破坏收集数据。本文基于地基激光雷达数据通过点云处理软件以人机交互的方式获取了26株落叶松样木的1266组枝条信息,包括着枝高度、弦长、枝长、着枝角度、基径和弓高。枝条可提取的最大相对着枝高度的平均值为0.83。在所提取的枝条因子中,提取精度依次为着枝高度＞弦长＞枝长＞基径(基径大于20 mm的枝条)＞弓高,将树冠分为4部分后分析发现,随着冠层高度的增加,枝条密度呈升高趋势,枝条提取率和提取精度呈下降趋势。此外,由于枝条基径提取精度较低,以弦长、着枝高度、胸径和树高为自变量构建基径预测模型。对不同基径的实测值、提取值与模型预测值对比分析发现,枝条基径的预测精度大于提取精度。对于造材来说,最有价值的部分是树木中下部,本方法能够较准确地提取树木胸径树高和相对着枝高度0.8以下的枝条属性信息,提供构建木材质量模型所需要的参数。     

Altered Growth and Wood Characteristics in Transgenic Hybrid Aspen Expressing Agrobacterium tumefaciens T-DNA Indoleacetic Acid-Biosynthetic Genes

A key regulator of cambial growth is the plant hormone indoleacetic acid (IAA). Here we report on altered wood characteristics and growth patterns in transgenic hybrid aspen (Populus tremula L. x Populus tremuloides Michx.) expressing Agrobacterium tumefaciens T-DNA IAA-biosynthetic iaaM and iaaH genes. Eighteen lines simultaneously expressing both genes were regenerated. Of these, four lines, verified to be transgenic by northern blot analysis, were selected and raised under controlled growth conditions. All four lines were affected in their growth patterns, including alterations in height and stem diameter growth, internode elongation, leaf enlargement, and degree of apical dominance. Two transgenic lines, showing the most distinct phenotypic deviation from the wild type, were characterized in more detail for free and conjugated IAA levels and for wood characteristics. Both lines showed an altered IAA balance, particularly in mature leaves and roots where IAA levels were elevated. They also exhibited changes in wood anatomy, most notably a reduction in vessel size, an increase in vessel density, and changes in ray development. Thus, the recent development of techniques for gene transfer to forest trees enabled us to investigate the influence of an altered IAA balance on xylem development in an intact experimental system. In addition, the results demonstrate the possibility of manipulating wood properties in a forest tree through controlled changes of IAA concentration and distribution.     

散孔材与环孔材树种枝干、叶水力学特性的比较研究

为揭示散孔材与环孔材树种树木水分生理特性的差异,选取了常见的3种散孔材落叶树种(毛白杨、法国梧桐和樱花)和3种环孔材落叶树种(刺槐、合欢和白蜡),研究了其枝干与叶水力学性质的差异及其协调性。结果表明:3种环孔材树种枝干横截面积基础上的最大比导水率(Ks-max)大于3种散孔材树种,但其木质部对空穴化的脆弱性(P50branch)高于散孔材树种,6种树木枝干的水分传输能力和抵抗空穴化能力之间存在一种相互制约的权衡关系。3种散孔材与3种环孔材树种的叶最大水力导度(Kl-max)和水力脆弱性(P50leaf)并无显著差异;对于3种散孔材树种,叶的水力脆弱性要高于枝干,但对3种环孔材树种而言,枝干的水力脆弱性要高于叶。6种树木枝干和叶的水力学性质(Kmax、P50)之间并无相关关系。这些结果表明:散孔材与环孔材树种的枝干水力学特性有明显差异,但叶水力学特性无差异;枝干与叶水力学性质之间是相互独立的。     

Effect of wood density and water permeability on wood decomposition rates of 32 Bornean rainforest trees

Aims A better understanding of wood litter decomposition is essential for predicting responses of forest ecosystems to global climate change. Recent studies suggest that chemical properties of wood litters, rather than physical ones such as wood density, are more important for interspecific differences in wood decomposition rates. However, empirical data are still limited, especially for tropical trees. In addition, decomposition rate of wood litter often varies with time, which makes interspecific comparison difficult. We studied the wood decomposition of 32 rainforest trees to elucidate (i) the degree of interspecific variation in wood decomposition rate of a given size and configuration and (ii) if initial wood density and water permeability are consistent predictors of the overall decomposition rate and its pattern over time.Methods A common garden decomposition experiment was conducted in a tropical rainforest in Malaysian Borneo for 32 native tree species. Small wood sticks were set on the forest floor and the weight loss was monitored monthly for 2.7 years.Important findings We found large variation in the wood decomposition rate (a 49-fold range), suggesting that we need to consider this variation when calculating community-level carbon dynamics of tropical rain forests. The physical traits of wood, i.e. wood density and water permeability, were related to wood decomposition rate and its pattern over time. Decomposition half-time related positively and negatively to initial wood density and water permeability, respectively. The time-dependent-rate model fitted better for 18 species (56% of the study species) that had higher water permeabilities than the others, suggesting that micelle porosity in wood relates to temporal changes in decomposition rate.     

Finding the optimal lengths for three branches at a junction

This paper presents an exact analytical solution to the problem of locating the junction point between three branches so that the sum of the total costs of the branches is minimized. When the cost per unit length of each branch is known the angles between each pair of branches can be deduced following reasoning first introduced to biology by Murray. Assuming the outer ends of each branch are fixed, the location of the junction and the length of each branch are then deduced using plane geometry and trigonometry. The model has applications in determining the optimal cost of a branch or branches at a junction. Comparing the optimal to the actual cost of a junction is a new way to compare cost models for goodness of fit to actual junction geometry. It is an unambiguous measure and is superior to comparing observed and optimal angles between each daughter and the parent branch. We present data for 199 junctions in the pulmonary arteries of two human lungs. For the branches at each junction we calculated the best fitting value of x from the relationship that flow ∞ (radius)^x. We found that the value of x determined whether a junction was best fitted by a surface, volume, drag or power minimization model. While economy of explanation casts doubt that four models operate simultaneously, we found that optimality may still operate, since the angle to the major daughter is less than the angle to the minor daughter. Perhaps optimality combined with a space filling branching pattern governs the branching geometry of the pulmonary artery.     

Mutual interaction of auxin and cytokinins in regulating correlative dominance

Correlative dominance requires correlative signals from a dominant to a dominated organ. Auxins, particularly IAA, and cytokinins are obviously important components of this correlative system. Using a vegetative pea shoot and a generative apple and tomato fruit system it can be demonstrated that dominant organs always export more IAA and have a higher ³H-IAA transport capacity and velocity compared to dominated organs. In both systems the dominant organ can be replaced by the application of auxin, e.g. NAA, which maintains the differences in IAA export. This is an indication that similar regulatory mechanisms control dominance in both of these diverse systems. The possibility of replacing a dominant organ by auxin also makes it unlikely that growth of that organ or allocation of nutrients regulates the correlative inhibition of the dominated organ.It is suggested that differences in IAA export from, and transport capacities of, dominant and dominated shoots, may be explained by a mechanism of auxin transport autoinhibition (ATA), whereby the earlier and stronger export of IAA from the dominant shoot inhibits auxin export from the dominated shoot at the point where the two auxin streams converge. This hypothesis was tested with explants of pea, apple and tomato. It was shown that the basal application of cold IAA significantly reduced endogenous as well as exogenous IAA transport through these explants.Since the reduced IAA transport of dominated organs was not followed by an accumulation of IAA in the auxin producing subtending organ, it was concluded that IAA biosynthesis was possibly reduced and/or IAA conjugation stimulated. This could have been one of the determinants of their growth inhibition. ATA might also explain how the unidirectional IAA signal may affect the growth rate of organs even lateral or acropetal to its transport pathway and thus polar IAA-transport becomes a ``multidirectional' signal. From the experiments demonstrated it seems that ATA is a sufficient mechanism to impose growth inhibition in the dominated organ, without the need of other regulators.However, to release dominated organs from dominance cessation of ATA may not be sufficient and cytokinins are obviously a powerful antagonist to auxins. Their repeated exogenous application turns dominated lateral buds into strongly growing organs which ultimately may even dominate the previously dominant apex. These lateral shoots finally gain a strong IAA export capacity and inhibit, by ATA, IAA export from the hitherto dominant apex.In other experiments it was shown that interruption of polar IAA transport leads to a strong increase in root derived cytokinins. This can largely be prevented, in a concentration dependent manner, by the application of auxin, indicating that basipolar auxin may control cytokinin production in the roots and its possible delivery to lateral buds. In turn, the increased delivery of cytokinins to the lateral buds promotes a strong increase in IAA production and export. Thus there is a strong mutual interaction between auxin production in the shoots and cytokinin production in the roots, which may be important in regulating the balance between root and shoot growth.     

植株大小、枝龄和环境共同驱动红松枝性状的变异

许多枝性状的变异受植株大小、枝龄或环境的影响, 但少有研究同时评估这些因素对枝性状种内变异的重要性。该研究以红松(Pinus koraiensis)为研究对象, 通过测定69株胸径(DBH) 0.3-100.0 cm范围内植株不同年龄枝的形态性状、化学性状和解剖性状, 探讨植株大小(DBH或树高)、枝龄与环境因素(光照强度、土壤养分及土壤含水率)对枝性状的影响。结果表明: (1) DBH与树高对枝性状的影响存在差异: 木质密度(WD)、木质部面积占比(R_XA)、韧皮部面积占比(R_PHA)及髓面积占比(R_PA)对DBH更敏感, 而树脂道总面积占比(R_RC)和枝氮含量(WN)受树高影响更大; (2)枝龄是导致红松枝性状种内变异的最主要因素, 植株大小次之, 而环境因素的影响最小; (3) WD、R_PHA与DBH显著正相关, R_PA与DBH显著负相关, R_RC、WN与树高显著正相关; 除WN外, 其余枝性状与枝龄均显著相关, 且随着树木生长, R_RC随枝龄增大而减小的速率加剧, 相反, R_PA随枝龄增大而减小的速率减缓。研究结果有助于了解局域尺度上枝性状种内变异的影响因素以及枝条应对环境变异的适应机制。     

SURFACE AREA RELATIONS OF WOODY PLANTS AND FOREST COMMUNITIES

Surface area of wood and bark is an important dimension of forests, with implications for respiration rate, energy exchange, and water and mineral budgets. Surface area of stem wood and bark can be estimated effectively from linear regressions on conic surface (one-half basal circumference times tree height) or from regressions of the logarithm of area on the logarithm of diameter at breast height. Branch surface can be estimated from a formula using branch basal diameter, length, and number of current twigs, and from logarithmic regressions of branch bark surface on basal diameter of branches and breast-height diameter of trees. In temperate deciduous forests several square meters of plant surface occur above each square meter of ground surface; these plant surfaces include 0.3–0.6 m² of stem bark, 1.2–2.2 m² of branch bark, and 3.0–6.0 m² of leaf blades. Branch bark surface increases more rapidly than leaf surface with increasing size of branches and trees. Growth and aging of trees, and maturation of forests, imply increasing ratios of bark (and wood) surface to the photosynthetic leaf surface which supports its growth and respiration.     

Branch development in Lupinus angustifolius L.#II. Relationship with endogenous ABA, IAA and cytokinins in axillary and main stem buds

The concentrations of indole-3-acetic acid (IAA), cytokinins (CK) and abscisic acid (ABA) were measured in buds of different regions (main stem and lateral branches) of Lupinus angustifolius L. (cv. Merrit) and at different stages in the development of branches. In lupin, branching patterns are the result of discrete regions of axillary branches (upper, middle and basal) which elongate at much different rates. Early in development only the main shoot elongates, followed usually by basal branch growth and then rapid upper branch growth. Branches in the middle of the main stem grow only weakly or fail to develop. Levels of IAA were generally high in the apical buds of slowly growing branches and low in buds from strongly growing branches, whereas CK levels showed the opposite relationship. CK:IAA ratio showed a closer relationship with the rate of growth of a particular branch better than the levels of either CK or IAA alone. During early stages of growth ABA concentration did not follow the rate of branch growth. However, later in development, where growth did not closely match the ratio of CK:IAA, ABA level showed a strong negative relationship with growth. A significant decrease in ABA was associated with continued strong growth of the main stem apex following a decline in CK:IAA ratio. Overall, the best relationship between the level of growth factors in apical buds and branching pattern in lupin was the ratio of CK:IAA, implying that high CK:IAA at a given bud would promote growth. ABA level appeared to play a secondary role, as a growth inhibitor.     

Origin of the biomechanical properties of wood related to the fine structure of the multi-layered cell wall

In this study, a basic model is introduced to describe the biomechanical properties of the wood from the viewpoint of the composite structure of its cell wall. First, the mechanical interaction between the cellulose microfibril (CMF) as a bundle framework and the lignin-hemicellulose as a matrix (MT) skeleton in the secondary wall is formulated based on "the two phase approximation." Thereafter, the origins of (1) tree growth stress, (2) shrinkage or swelling anisotropy of the wood, and (3) moisture dependency of the Young's modulus of wood along the grain were simulated using the newly introduced model. Through the model formulation; (1) the behavior of the cellulose microfibril (CMF) and the matrix substance (MT) during cell wall maturation was estimated; (2) the moisture reactivity of each cell wall constituent was investigated; and (3) a realistic model of the fine composite structure of the matured cell wall was proposed. Thus, it is expected that the fine structure and internal property of each cell wall constituent can be estimated through the analyses of the macroscopic behaviors of wood based on the two phase approximation.     

尾叶桉人工林种群密度的研究

探讨了5.6年生尾叶桉种群密度与冠幅、胸径、树高、立木单株材积、林分蓄积量、木材性质及保存率等的作用规律和相关模型.结果表明,密度对胸径、立木单株材积、冠幅及枝下高的影响达到极显著水平;对蓄积量、木材纤维宽度的影响达显著水平;对树高、木材气干密度和木材纤维长度虽有一定的影响,但不显著.其中,密度与蓄积量、枝下高、木材纤维宽度呈正相关关系;而与胸径、立木单株材积、冠幅呈负相关关系.此外,尾叶桉具有较宽的合理密度范围;作为短周期浆纸林,其最佳密度应确定为2000株·hm_-2.     

尾叶桉人工林种群密度的研究

探讨了５．６年生尾叶桉种群密度与冠幅、胸径、树高、立木单株材积、林分蓄积量、木材性质及保存率等的作用规律和相关模型。结果表明,密度对胸径、立木单株材积、冠幅及枝下高的影响达到极显著水平;对蓄积量、木材纤维宽度的影响达显著水平;对树高、木材气干密度和木材纤维长度虽有一定的影响,但不显著。其中,密度与蓄积量、枝下高、木材纤维宽度呈正相关关系;而与胸径、立林单株材积、冠幅呈负相关关系。此外,尾叶桉具有较