Application of the Functional-Structural Tree Model LIGNUM to Sugar Maple Saplings (Acer saccharum Marsh) Growing in Forest Gaps

LIGNUM is a functional-structural model that represents a treeusing four modelling units which closely resemble the real structureof trees: tree segments, tree axes, branching points and buds.Metabolic processes are explicitly related to the structuralunits in which they take place. Here we adapt earlier versionsof LIGNUM designed to model growth of conifers for use withbroad-leaved trees. Two primary changes are involved. First,the tree segment for broad-leaved trees consists of enclosedcylinders of heartwood, sapwood and bark. Leaves consistingof petioles and blades are attached to the segments. Secondly,axillary buds and rules governing their dormancy are includedin the model. This modified version of LIGNUM is used to simulatethe growth and form of sugar maple saplings in forest gaps.The annual growth of the model tree is driven by net productionafter respiration losses are taken into account. The productionrate of each leaf depends on the amount of photosyntheticallyactive radiation it receives. The radiation regime is trackedexplicitly in different parts of the tree crown using a modelof mutual shading of the leaves. Forest gaps are representedby changing the radiation intensity in different parts of themodel sky. This version of LIGNUM modified for use with broad-leaf,deciduous trees and parameterized for sugar maple, yields goodsimulations of growth and form in saplings from different forestgap environments. Copyright 2001 Annals of Botany Company LIGNUM, functional-structural tree model, tree architecture, sugar maple, modelling growth and form     

Branching Principles Governing the Architecture of Cornus kousa (Cornaceae)

The complex structure of the crown of Cornus kousa, generallyfive-forked in vegetative branching and two-forked in reproductivebranching, is analysed quantitatively and described by two basicbranching principles: decussate phyllotaxy and the resettingrule for planes of branching. Most Cornus species have opposite,decussate phyllotaxis. The leaf pair (with axillary buds) definesthe branching plane of a node. Because of regular phyllotaxis,the fundamental branching pattern is that every branching planealong an axis is perpendicular to the preceding one. However,the first node of a lateral horizontal shoot always has a horizontalbranching plane; we term this the resetting rule. We observedthat resetting occurs when the first nodes initiated in thevertical plane are repositioned by a twisting of their firstinternodes. All later nodes alternate directions, i.e. showusual decussate alternation. Foliage leaf nodes usually producethree-forked branchings. When vegetative winter buds are formed,a foliar node and adjacent scale leaf node produce a five-forkedbranching. When reproductive winter buds with a terminal inflorescenceare formed, the last foliar node and two adjacent scale leafnodes can produce a variety of branchings but usually producean equal two-forked branching. To understand better the architecturein C. kousa, we contrast it with C. capitata which does notproduce buds with scale leaves and whose vegetative nodes areclearly separated. Copyright 1999 Annals of Botany Company Branching pattern, Cornaceae, Cornus kousa, decussate branching, dogwood, Japanese strawberry tree, tree architecture, tree geometry.     

Tree roots as self-similar branching structures: axis differentiation and segment tapering in coarse roots of three boreal forest tree species

We applied a fractal root model to the 3D architecture of the coarse root systems of Betula pendula Roth, Picea abies (L.) H. Karst., and Pinus sylvestris L. in mixed boreal forests. Our dataset consisted of 60 root systems excavated in five different mixed forest stands. We analyzed the variability of the model parameters with respect to species, site type, and different root axes. According to our results, the cross-sectional area of root segments (i.e. second power of diameter) was a suitable variable for analyzing the values of parameters of the fractal model. The parameter values varied with generation and order of root segments; the roots thus did not follow the simple fractal branching. The variation of parameters along the root axes showed the existence of a zone of rapid tapering in all tree species. The model was, with parameter values analyzed from the data, moderately capable of accounting for the main coarse root characteristics. It was important for model predictions to take into account the tapering of root segments. We conclude that, in boreal forests, tree root systems are the output of the axis-specific morphogenetic branching rules and functional adaptation to spatial heterogeneity in the soil.     

Growth Analysis of Perennial Plants: The Relative Production Rate and its Yield Components

A new formulation for plant growth analysis, called the relativeproduction rate, is proposed for use in the study of perennialplants. The new measure and its yield components are similarto current growth analysis, but use the annual increment ofgrowth, rather than total accumulated growth, as the basis forassessing performance. The relative production rate removesaccumulated past growth of the perennating structure, a majordeterminant of relative growth rate, and acts independentlyof tree size as a measure of the vigour of growth. The multiplicativeyield components of this relative production rate also providebetter insight into the current morphological and physiologicalfeatures of the tree than the standard growth analysis formulations.Examples are given using Douglas-fir (Pseudotsuga menziesii(Mirb.) Franco) and red pine (Pinus resinosa Ait.) plantationsto illustrate the methodology proposed. Pseudotsuga menziesii (Mirb.) Franco, Pinus resinosa Ait., Douglas-fir, red pine, growth analysis, relative production rate, tree development, competition     

Quantitative characterization of clumping in Scots pine crowns

Background and Aims

Proper characterization of the clumped structure of forests is needed for calculation of the absorbed radiation and photosynthetic production by a canopy. This study examined the dependency of crown-level clumping on tree size and growth conditions in Scots pine (Pinus sylvestris), and determined the ability of statistical canopy radiation models to quantify the degree of self-shading within crowns as a result of the clumping effect.

Methods

Twelve 3-D Scots pine trees were generated using an application of the LIGNUM model, and the crown-level clumping as quantified by the crown silhouette to total needle area ratio (STAR_crown) was calculated. The results were compared with those produced by the stochastic approach of modelling tree crowns as geometric shapes filled with a random medium.

Key Results

Crown clumping was independent of tree height, needle area and growth conditions. The results supported the capability of the stochastic approach in characterizing clumping in crowns given that the outer shell of the tree crown is well represented.

Conclusions

Variation in the whole-stand clumping index is induced by differences in the spatial pattern of trees as a function of, for example, stand age rather than by changes in the degree of self-shading within individual crowns as they grow bigger.     

Persistent Effects of Short-term, High Exposure to Chlorine Gas on Physiology and Growth of Pinus ponderosa andPseudotsuga menziesii

Following a single acute exposure to chlorine gas, persistenteffects on epicuticular waxes, cuticular transpiration, treegrowth and mortality were studied in foliage of Pinus ponderosaand Pseudotsuga menziesii for three growing seasons. Chlorinegas exposure caused foliar injury to both exposed foliage andfoliage that flushed after exposure (P < 0.05). The tendencyto form films of water rather than droplets was greater in directlyexposed foliage (P < 0.001). Rates of cuticular transpirationwere higher for directly and indirectly exposed foliage of Pinusponderosa up to 1 year after exposure and up to 6 months afterexposure for directly exposed Pseudotsuga menziesii(P < 0.001),after which P. menziesii needles defoliated. Total water content(TWC) and relative water content were significantly correlatedwith foliar injury (P < 0.05). TWC was lower for directlyexposed foliage up to 1 year after exposure (P < 0.001).There was no persistent negative effect on F_v/F_m ratios after1 year. Exposure to chlorine gas did not affect needle lengthor annual shoot increment growth, but exposure was correlatedwith increased bud production. Needle longevity of foliage thatflushed 2 months after exposure was reduced significantly (P< 0.001). Annual stem increment growth for both species decreasedover at least three growing seasons following chlorine gas exposure(P < 0.001), and depended on distance from the spill site.Cone production was lower for exposed Pinus ponderosa treescompared to controls (P < 0.05), and tree mortality was higherwithin approx. 50 m of the release site forPseudotsuga menziesii. Growth responses for both conifers agreed well with predictedpatterns of carbon allocation after defoliation caused by chlorinegas exposure. Copyright 2001 Annals of Botany Company Pinus ponderosa, Pseudotsuga menziesii, conifers, chlorine gas, leaf wettability, cuticular transpiration, water relations, growth, mortality     

Age-related decline in stand productivity: the role of structural acclimation under hydraulic constraints

The decline in above-ground net primary productivity (P_a) that is usually observed in forest stands has been variously attributed to respiration, nutrient or hydraulic limitations. A novel model is proposed to explain the phenomenon and the co-occurring changes in the balance between foliage, conducting sapwood and fine roots. The model is based on the hypothesis that a functional homeostasis in water transport is maintained irrespective of age: hydraulic resistances through the plant must be finely tuned to transpiration rates so as to avoid extremely negative water potentials that could result in diffuse xylem embolism and foliage dieback, in agreement with experimental evidence. As the plant grows taller, allocation is predicted to shift from foliage to transport tissues, most notably to fine roots. Higher respiration and fine root turnover would result in the observed decline in P_a. The predictions of the model have been compared with experimental data from a chronosequence of Pinus sylvestris stands. The observed reduction in P_a is conveniently explained by concurrent modifications in leaf area index and plant structure. Changes in allometry and shoot hydraulic conductance with age are successfully predicted by the principle of functional homeostasis.     

Dry-matter Production by Pinus sylvestris L.

This paper contains the data for the gross weight and relativeproportions of canopy, bole, and roots in a series of twelveeven-aged forest stands. From these it is possible to describe the growth curves of treesof Pinus sylvestris in Britain under the observed conditionsand the production of dry matter per unit area of such standsof trees. The following general results are given: regressionsrelating weight to bole, height, and diameter; the relativegrowth of leaves, roots, and bole; annual increments of drymatter; relative growth-rates per unit leaf area or per unitdry weight per annum. Some comparative data are given for Pinus nigra (one plantation)which is more productive than Pinus sylvestris.     

The Effects of Simazine, Applied for Weed Control, on the Mycorrhizal Development of Pinus Seedlings

The herbicide Simazine is widely used in forestry nurseries.Its effect on the incidence of ectomycorrhizae on pine seedlingroot systems was investigated. Under field (nursery) conditions,at routine rates of application, Simazine does not inhibit mycorrhizaldevelopment in Pinus sylvestris and P. nigra, and it may, undersome conditions, actually enhance mycorrhizal formation. Sucha stimulatory effect was also obtained with pot-grown seedlingstreated with the herbicide. It is suggested that soil type andclimatic factors might be involved in determining the responseto Simazine. The mode of stimulation of mycorrhizal developmentby this herbicide is not understood, but it is not via suppressionof weed growth. Pinus sylvestris, Pinus nigra, pine seedlings, ectomycorrhizae, Simazine     

河北坝上樟子松人工林径向生长及其对气候因素的响应

利用树木年轮学方法测定了河北坝上地区樟子松逐年树轮宽度和胸高断面积增量,建立了樟子松树轮宽度差值年表并分析其对气候因素的响应以及生态弹性,为河北坝上地区利用樟子松开展退化杨树防护林更新建设的适宜性提供理论参考。研究结果表明,河北坝上地区樟子松树龄在28 a左右,其生长阶段可划分为快速生长期(0—10 a),生长下降期(11—20 a)和生长平稳期(21—28 a);樟子松树轮宽度在1992—2000年期间为快速增长期((4.49±1.15)mm/a),在2001—2010年期间呈现下降趋势((3.62±1.39)mm/a),而在2011—2019年期间樟子松生长呈平稳特征((2.21±0.68)mm/a),约为快速增长期增长速度的一半;樟子松BAI在1992—2000年间一直呈上升趋势((5.09±2.26)cm~2/a),2001—2019年期间呈平稳特征((10.46±0.67)cm~2/a),表现出稳定且持续的生长能力。樟子松差值年表与气候因素的相关性显示：樟子松径向生长主要与上一年生长季末期(9、10月)、休眠期(12月)及当年生长季(7月)平均温度和最高温度呈显著负相关,其中...     

A Model of Grass Growth

A model of grass growth is constructed and applied to above-grounddry-weight data on a well-fertilized and irrigated grass cropgrown at Hurley. The processes of light interception, photosynthesis,respiration, growth and senescence are represented in the model.Temperature, daily light receipt and daylength are the environmentinput variables required. The model is able to simulate thetwo limiting cases of exponential growth, and a steady-statecrop which is in dynamic equilibrium without net growth. Themodel fits the data well, and provides a simple framework forthe analysis of the growth of grass. Lolium perenne L., perennial ryegrass, light interception, photosynthesis, growth, respiration, senescence, mathematical model     

Shoot Elongation in Scots Pine: Diurnal Variations and Response to Temperature

Kanninen, M. 1985. Shoot elongation in Scots pine: Diurnal variationsand response to temperature.—J. exp. Bot. 36: 1760–1770. Time series analysis is used to study the diurnal variationsin the rate of shoot elongation of Scots pine (Pinus sylvestrisL.) seedlings, and its response to temperature. The shoot elongation of three 5-year-old Scots pine seedlingswas measured at 2 h intervals over a 12 d period in June 1979.Ambient temperature, measured with a thermocouple, was continuouslyrecorded during the same period. The measurements were carriedout in the field. Time series analysis of the data was carried out by estimatedauto-correlation and partial auto-correlation functions. A multiplicative‘seasonal’ model was applied to both the input andoutput series to ‘prewhiten’ the data set. Serialcross-correlation analysis of the prewhitened series was usedto identify the structure of the transfer function model betweentemperature and growth rate. There was a phase shift between the air temperature and shootextension. The cross-correlation function peaked at 2 h lagvalue. The influence of temperature on the growth rate was adequatelydescribed by a first-order transfer function model. The characteristicfeature of the model was a time constant of 3.3 h and a basetemperature of 6.3 °C for the response of shoot elongationto temperature. Key words: Scots pine, Pinus sylvestris L., shoot elongation, temperature response, time-series analysis     

The Correlation between Plant Growth and Intercepted Radiation: An Interpretation in Terms of Optimal Plant Nitrogen Content

Photosynthesis of leaves is commonly observed to have a saturatingresponse to increases in their nitrogen (N) content, while theresponse of plant maintenance respiration is more nearly linearover the normal range of tissue N contents. Hence, for a givenamount of foliage, net primary productivity (NPP) may have amaximum value with respect to variations in plant N content.Using a simple analytically-solvable model ofNPP, this ideais formulated and its broad implications for plant growth areexplored at the scale of a closed stand of vegetation. The maximum-NPPhypothesis implies thatNPP is proportional to intercepted radiation,as commonly observed. The light utilization coefficient (     

Detection of Intracellular Bacteria in the Buds of Scotch Pine (Pinus sylvestris L.) by In Situ Hybridization

Bacterial isolates were obtained from pine (Pinus sylvestris L.) tissue cultures and identified as Methylobacterium extorquens and Pseudomonas synxantha. The existence of bacteria in pine buds was investigated by 16S rRNA in situ hybridization. Bacteria inhabited the buds of every tree examined, primarily colonizing the cells of scale primordia and resin ducts.     

Stem Diameter Growth of Scots Pine Trees after Increased Mechanical Load in the Crown during Dormancy and (or) Growth

A field experiment with a 2 x2 factorial block design (W_xS_x)was conducted in northern Sweden where the mechanical loadsin the crowns of sixteen 2.5m high Scots pine (Pinus sylvestrisL.)trees were increased during one winter (W₁, dormant period)and (or) summer (S₁, growth period). Trees treated were loadedwith five 2kg bags hung over mid-crown branches close to thestem, i.e. 10kg per tree. After treatment, all trees were leftto grow untreated for one additional year. Trees were then cutat ground level and annual ring widths for the last 5 yearswere measured on stem discs taken at 1, 5, 10, 15, 20, 30 and50% of tree height. Differences between treatments were analysedwith two-way factorial ANOVA. Accumulated treatment responsewas positive for winter loading (W₁S_x) at all levels, and statisticallysignificant at 1, 15 and 20% of tree height. Summer loading(W_xS₁) had positive effects at the lowest and middle parts ofthe stem, and negative in between. No statistically significanttwo-way interaction (W xS) was observed. Results support thehypothesis that Scots pine trees can retain information aboutmechanical forces acting on their stems during winter, and respondto this during the following growth period. The results alsosuggest that stem form of trees in boreal forests may be stronglyaffected by winter conditions. Stem form; mechanical perturbation; Scots pine; Pinus sylvestris; dendrometer; diameter; growth; dormancy; thigmomorphogenesis; wind; sway     

Modelling Woody Plant Tissues5

An electrical impedance spectrum (20 Hz to 1 MHz) was measuredin both the bark and wood of current-year and one-year-old Scotspine (Pinus sylvestris L.) shoots. The measured impedance spectrawere analysed in reference to two lumped circuits and a distributedcircuit. It was found that neither of the two lumped circuitsfit the data from bark or wood as well as the distributed model.The lumped (double-shell) model fit fairly well for bark, butpoorly for wood. It is proposed that the good fit of the distributedmodel and the poorer fit of the lumped models are due to a widerange of cell sizes in the bark and wood. The distributed circuitused in the present study may be useful for describing differentiatedplant tissues for physiological studies. Key words: Bark, electrical impedance, equivalent circuit, Pinus sylvestris L., wood     

Bud Structure in Relation to Shoot Morphology and Position on the Vegetative Annual Shoots of Juglans regia L. (Juglandaceae)

The length and basal diameter of all lateral and terminal budsof vegetative annual shoots of 7-year-oldJuglans regia treeswere measured. All buds were dissected and numbers of cataphylls,embryonic leaves and leaf primordia were recorded. Each axillarybud was ranked according to the position of its associated leaffrom the apex to the base of its parent shoot. Bud size andcontent were analysed in relation to bud position and were comparedwith the size and number of leaves of shoots in equivalent positionswhich extended during the following growing season. Length andbasal diameter of axillary buds varied according to their positionon the parent shoot. Terminal buds contained more embryonicleaves than any axillary bud. The number of leaves was smallerfor apical and basal axillary buds than for buds in intermediatepositions on the parent shoot only. All new extended shootswere entirely preformed in the buds that gave rise to them.Lateral shoots were formed in the median part of the parentshoot. These lateral shoots derived from buds which were largerthan both apical and basal ones. Copyright 2001 Annals of BotanyCompany Juglans regia L., Persian walnut tree, branching pattern, preformation, bud content, shoot morphology     

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°。     

Photoregulation of growth and branching of plum shoots: Physiological action of two photosystems

Summary Plum shoot proliferation was investigated in terms of two distinct processes: axillary bud differentiation and axillary shoot development. Results showed that light quality influenced bud differentiation and interacted with apical dominance in determining shoot outgrowth, resulting in a differentiated structure of shoot clusters and type of branching. Results suggested that blue light, acting through its photoreceptor, increased the number of axillary buds differentiated from apical meristem, but did not remove the apical dominance. Red light removed apical dominance, while reducing the formation of axillary buds; both events appeared to be dependent on the putative amount of phytochrome active form, and independent of light photon fluence rate. On the contrary, blue light action appeared to be dependent on photon fluence rate. In addition, apparent blue-red interactions related to photomorphogenic events fit an antagonistic model for branching regulated by light via cryptochrome and phytochrome photoreceptors. Our results show that the dynamics of shoot cluster development is the product of two events: the formation of new axillary buds and their release from apical dominance.     

Modelling and Measuring Transpiration from Scots Pine with Increased Temperature and Carbon Dioxide Enrichment

Starting in 1996, individual trees of Scots pine (Pinus sylvestrisL.) aged 30 years, were grown in closed-top chambers and exposedto either normal ambient conditions (CON), elevated CO₂(approx.700 µmol mol^-1; Elev. C), elevated temperature (approx.2 °C and approx. 6 °C above the outside ambient temperatureduring the ‘growing season’ and ‘off season’,respectively; Elev. T) or a combination of elevated CO₂and warmertemperature (Elev. CT). Sap flow was monitored simultaneouslyby the constant-power heat balance method in a total of 16 trees,four for each treatment, over a 32 d period in summer 1998 (afterthe completion of needle expansion and branch elongation). Toquantify the contributions of crown and physical environmentalvariables to total crown transpiration, a ‘sun/shade model’was developed and used to partition the changes in transpirationto different sources. The results of the sap flow measurementsindicate that (1) total daily sap flow (E_tree.d) varied from0.15–3.41 kg per tree; (2) the treatment effect on E_tree.ddependedgreatly on the weather conditions; (3) the cumulative E_tree.dforthe 32 d dropped significantly by 22% relative to CON (P =0.038)under Elev. C and increased significantly by 21% (P =0.043)and 16% (P =0.048) under Elev. T and Elev. CT, respectively.In general, the modelled transpiration gave good agreement withthe sap flow results. The model computations showed that, ona typical sunny day in summer, the effect of treatment on crownstomatal conductance was responsible for approx. 80% of thechange inE_tree.d , while the increase in needle area and theeffect on total radiation absorption contributed only a smallpercentage. Furthermore, sunlit needles were responsible forover 60% of change in transpiration. The effect of the treatmentson E_tree.dwas larger at high temperature and vapour pressuredeficit but was not sensitive to incident daily radiation. Copyright2000 Annals of Botany Company Transpiration model, sap flow, CO₂and temperature elevation, environment-controlled chamber, Pinus sylvestris L.