Uniform versus Asymmetric Shading Mediates Crown Recession in Conifers

In this study we explore the impact of asymmetrical vs. uniform crown shading on the mortality and growth of upper and lower branches within tree crowns, for two conifer species: shade intolerant lodgepole pine (Pinus contorta) and shade tolerant white spruce (Picea glauca). We also explore xylem hydraulics, foliar nutrition, and carbohydrate status as drivers for growth and expansion of the lower and upper branches in various types of shading. This study was conducted over a two-year period across 10 regenerating forest sites dominated by lodgepole pine and white spruce, in the lower foothills of Alberta, Canada. Trees were assigned to one of four shading treatments: (1), complete uniform shading of the entire tree, (2) light asymmetric shading where the lower 1/4–1/3 of the tree crown was shaded, (3) heavy asymmetric shading as in (2) except with greater light reduction and (4) control in which no artificial shading occurred and most of the entire crown was exposed to full light. Asymmetrical shading of only the lower crown had a larger negative impact on the bud expansion and growth than did uniform shading, and the effect was stronger in pine relative to spruce. In addition, lower branches in pine also had lower carbon reserves, and reduced xylem-area specific conductivity compared to spruce. For both species, but particularly the pine, the needles of lower branches tended to store less C than upper branches in the asymmetric shade, which could suggest a movement of reserves away from the lower branches. The implications of these findings correspond with the inherent shade tolerance and self-pruning behavior of these conifers and supports a carbon based mechanism for branch mortality – mediated by an asymmetry in light exposure of the crown.     

Understanding crown shyness from a 3-D perspective

Background and AimsCrown shyness describes the phenomenon whereby tree crowns avoid growing into each other, producing a puzzle-like pattern of complementary tree crowns in the canopy. Previous studies found that tree slenderness plays a role in the development of crown shyness. Attempts to quantify crown shyness have largely been confined to 2-D approaches. This study aimed to expand the current set of metrics for crown shyness by quantifying the characteristic of 3-D surface complementarity between trees displaying crown shyness, using LiDAR-derived tree point clouds. Subsequently, the relationship between crown surface complementarity and slenderness of trees was assessed.MethodsFourteen trees were scanned using a laser scanning device. Individual tree points clouds were extracted semi-automatically and manually corrected where needed. A metric that quantifies the surface complementarity (S_c) of a pair of protein molecules is applied to point clouds of pairs of adjacent trees. Then 3-D tree crown surfaces were generated from point clouds by computing their α shapes.Key ResultsTree pairs that were visually determined to have overlapping crowns scored significantly lower S_c values than pairs that did not overlap (n = 14, P < 0.01). Furthermore, average slenderness of pairs of trees correlated positively with their S_c score (R² = 0.484, P < 0.01), showing agreement with previous studies on crown shyness.ConclusionsThe characteristic of crown surface complementarity present in trees displaying crown shyness was succesfully quantified using a 3-D surface complementarity metric adopted from molecular biology. Crown surface complementarity showed a positive relationship to tree slenderness, similar to other metrics used for measuring crown shyness. The 3-D metric developed in this study revealed how trees adapt the shape of their crowns to those of adjacent trees and how this is linked to the slenderness of the trees.     

Shoot damage by Tomicus sp. (Coleoptera: Scolytidae) and effect on Pinus yunnanensis resistance to subsequent reproductive attacks in the stem

Abstract 1 In South‐western China, Yunnan pines Pinus yunnanensis, suffer considerable damage from an undescribed Tomicus sp. previously thought to be T. piniperda. 2 To assess the effect of shoot maturation feeding (during which an aggregation process appears to occur) on host resistance to attacks on the bole, the relationships between shoot damage, bole attack density and tree survival were studied. 3 Attack distribution in the crown and in the stem did not vary between killed and surviving trees, indicating that mortality is determined by the quantity of attacks. 4 The level of shoot damage and bole attack density were positively and linearly correlated. This can be explained by the fact that bole attacks are caused by beetles coming from the crown of the same tree. 5 A critical threshold of bole attack density (around 80 attacks/m²) above which trees die was observed. However, because attacks continue after this threshold is reached, the density of failed attacks on the killed trees was used as an estimator of the threshold density. It decreased when shoot damage increased. 6 The existence of a critical threshold of shoot damage (60% damaged shoots) was also demonstrated. Above this threshold, stem attack density was always sufficiently high to kill trees. 7 The results emphasize that concentration of shoot attacks is the main reason for the extensive tree damage observed in China. 8 A model of relationships between shoot and stem attacks is proposed, suggesting that management to reduce shoot attacks would protect trees from dying by both decreasing the number of bole attacks and raising the threshold for successful attack density on the bole to levels that could not be attained.     

Effects of branch position on water relations and gas exchange of European larch trees in an alpine community

Water relations and gas exchange were studied in the crowns of small European larch (Larix decidua Mill.) trees with respect to branch position. The upper-crown branches showed significantly higher branch sap flux rate (F _la) and branch conductance (g _b) compared to the lower crown (P<0.001). Values of leaf conductance (g _l), transpiration rate (E) and net photosynthesis (A), averaged for different ranges of atmospheric vapour pressure deficit (VPD), were also higher in the upper crown position. We suppose that the up to 2.6-fold smaller soil-to-leaf hydraulic conductance observed in the lower branches (P<0.001, compared to upper branches) could contribute to the decreased values of F _la, g _b, g _l, and E in the lower crown position. Variation in tracheid lumen diameter with respect to crown position (P<0.001) supported the hypothesis that branches growing at the crown base are hydraulically more constrained than branches located at the top of the tree. Leaf area to sapwood area ratio (A _la/A _sa) exhibited 1.4 times smaller values in lower crown (P<0.01), however, this could not compensate the effect of decreased hydraulic conductivity of the lower-crown branches.     

Photosynthesis and nitrogen allocation in needles in the sun and shade crowns of hybrid larch saplings: effect of nitrogen application

We studied the effects of applying 50 kg(N) ha^?1 year^?1 of nitrogen (N) on needle photosynthesis, N allocation and nutrient content in the sun- and shade crowns of the hybrid larch F₁ (Larix gmelinii var. japonica × L. kaempferi). The light-saturated net photosynthetic rate (P _Nmax) was not significantly affected by N application or crown position, although the contents of N, P, K, and chlorophyll (Chl), and the maximum rates of carboxylation and electron transport were lower in needles of the shade crown than of the sun crown. This difference was mainly due to an increase in the intercellular CO₂ concentration (C _i) in the needles of the shade crown. Analysis of N allocation in photosynthetic systems revealed that more N was allocated to functions related to electron transport and ribulose-1,5-bisphosphate (RuBP) regeneration in needles of the shade crown. N allocation in needles of the hybrid larch F₁ was regulated mainly by the light conditions, rather than by N application     

落叶松人工林生物量模型研究

以不同年龄、不同密度的落叶松（Larix olgensis）人工林为研究对象,基于19块标准地95株标准木的树干解析、枝解析的生物量数据,研究不同大小树木因子（胸径、树高、冠幅等）与单木各分量（树干、枝、叶）生物量之间的关系,应用统计分析软件建立落叶松单木各部分生物量的回归模型。利用单木各部分生物量回归模型方程估测落叶松人工林各林分的总生物量,并分析了不同年龄及林分密度下林分生物量的变化规律：林分的生物量随年龄的增加而不断增长,树干的生物量的比例是最大的,同时也是随着年龄的增长而不断的增加,而树枝和树叶的生物量的比例比较小,林分的生物量随林分密度的增加而不断增加。最后建立林分生物量模型,为落叶松人工林的研究提供基础资料,为了解落叶松人工林的生产力,对其进行合理经营提供科学依据。     

Nonlinear mixed-effects branch diameter and length models for natural Dahurian larch (<Emphasis Type="Italic">Larix gmelini</Emphasis>) forest in northeast China

Key message

We developed the generalized branch diameter and length models using the multi-level nonlinear mixed-effects techniques for the natural Dahurian larch ( Larix gmelini ) forest in northeast China.

Abstract

Dahurian larch (Larix gmelini) is the most commercially cultivated timber species in northeastern China due to its ecological prevalence and its superior wood attribute. However, its timber quality was largely driven by the crown architecture, i.e., the number, size and distribution of branches. The majority of branch-level models in the literature are focused on planted forests, which have substantially different crown architecture than that grown in natural mixed forests. Therefore, the goal of this investigation was to develop branch diameter and length models for Dahurian larch that are grown in natural mixed forests. A multi-level nonlinear mixed-effects model technique, including the fixed-effects, random-effects, variance functions and correlation structures, was employed to develop the branch growth models. The results suggested that the cumulative branch diameter and length were both increased with the increases of branch depth into the crown. Diameter at breast height (DBH) had significant positive influences on the branch size; however, tree height (HT) produced negative influences on the branch size, i.e., larger DBH and smaller HT could lead to larger branch size. Model fitting and validation results confirmed that we should avoid developing over-complex models from the perspective of application. As for the branch diameter and length models in our study, addressing the stand and tree level effects as random component were quite reliable and accurate for predicting the branch growth process of Dahurian larch in northeastern China.

     

Plastic Response of Crown Architecture to Crowding in Understorey Trees of Two Co-dominating Conifers

Crown architecture and growth rate of trunk height, trunk diameterand lateral branches of understorey trees (5–10m tall)were compared between two co-dominating conifers,Abies sachalinensisandPiceaglehnii, in relation to the index of local crowding intensity,W,represented as a function of density, distance and basal areaof taller neighbours. For the two conifers, the growth of trunkheight and diameter was decreased and crowns became flat withincreasingW, keeping crown projection area. Self-pruning oflower branches was more intense inAbiesthan inPiceain crowdedconditions, while both conifers showed similar crown forms inless crowded conditions. These results suggest that the growthin lateral branches exceeded that in height in crowded conditions,especially inAbies. Tree age of both conifers increased withincreasingW, resulting from the low growth rate in crowded conditions.The age of the longest and lowest branch ofPicea, up to 150years, was positively correlated with the tree age ranging from70 to 250 years, whereas that ofAbieswas constant at around30 years irrespective of tree age varying from 40 to 140 years.This result agrees with the observation that agedAbieshad moreflat-shaped crowns than in agedPiceain crowded conditions. Theseresults suggest that each conifer adapted to crowding in differentways: high elongation of branches with high turnover rate forAbiesandviceversaforPicea. Abies; crown form; neighbourhood interference; Picea; plasticity     

思茅松天然林树冠结构模型

以云南省普洱市思茅区思茅松天然林为研究对象,采用枝解析调查了34株思茅松样木的树冠数据,分析了一级枝枝长、枝径、着枝角度、弦长和树冠半径5个树冠形状变量的变化规律,分别构建其预估模型;分析了树冠结构变化,分别构建了一级枝轮枝高度预估模型、一级枝枝条数量预估模型和一级枝枝条数量累积预估模型,并采用独立样本进行模型统计精度检验。结果表明:8个预估模型的预测效果良好,精度达到91%以上,尤其是一级枝着枝角度模型和一级枝轮枝高度模型预测精度达到97%以上。研究结果合理准确描述思茅松树冠结构的变化,为思茅松天然林的经营管理提供科学依据。     

沙地云杉构件动态与结构

研究了沙地云杉的构件种群。结果表明：（1）沙地云杉根系生长迅速;（2）针叶寿命随树龄而变化;（3）枝条数量与枝条年龄存在如下关系：B40＝2682.5e-0.6958A,B200=5807.5e-1.3112A;（4）雄花／雌花随树龄迅速上升;（5）不同层次和方向上球果数量存在显著差异;（6）东方向的冠半径最长,西方向最短,枝条数量、长度,小枝数量随树龄而增加,小枝长度随树龄而缩短。结果表明,沙地云杉能够适就干旱的生态环境,其构件动态与结构是长期的自然选择与适应的结果。     

ARCHITECTURE AND CROWN GEOMETRY IN TABEBUIA ROSEA (BIGNONIACEAE)

In the tropical tree Tabebuia rosea, seedlings form an erect unbranched stem with rhythmic growth. Three distinct and predictable geometrical stages are then recognized in crown development. Sympodial branching begins with the arrest of the terminal bud of the trunk and symmetric outgrowth of a pair of subtending lateral buds. During an intermediate phase, branching becomes asymmetric at about Order 5. At each sympodial bifurcation there is differentiation between vigorous, relatively straight main branches (leaders) and less vigorous laterals forming regular pseudomonopodial branch complexes, which collectively constitute the cup-shaped crown. Finally, dormant lateral buds in the lowest bifurcation of the trunk are released and reiterate the original crown form. Ultimately an erect, apparently monopodial tree is formed by a set of superimposed cup-shaped crowns. Crown development of Tabebuia is unique because it involves predictable ontogenic changes in branching patterns. Crowns of open-grown Tabebuia consist of relatively few, wide branch tiers, crowns of forest grown trees are tall and narrow. Analysis of the adaptive geometry of wide vs. narrow crowns through computer simulation illustrates the precise cost of mechanical support for terminal leaf rosettes at successive developmental phases and suggests that tall, erect, narrow, and multi-tiered crowns are more efficient than wide open crowns.     

The tree crown distribution of hemlock woolly adelgid, Adelges tsugae (Hem., Adelgidae) from randomized branch sampling

Abstract:  An exotic insect, the hemlock woolly adelgid, Adelges tsugae Annand (Hem., Adelgidae), is spreading through eastern North America, killing hemlock trees [ Tsuga canadensis (L.) Carrière], and thereby impoverishing ecosystems. Adelges tsugae , like many alien invasive insects, is difficult to monitor or sample in the forest. Monitoring of A. tsugae has been hampered by lack of information about its distribution within tree crowns. In order to assist future monitoring and biocontrol of A. tsugae , this study investigates the crown distribution of A. tsugae by sampling from the entire height of mature hemlock trees in a forest with an established infestation. In addition to A. tsugae , sampling includes scale insects, which are another group of important pests on hemlock trees. This study demonstrates the utility of a randomized branch sampling (RBS) plan for monitoring both invasive insects as well as native insects that are difficult to sample. Results from the RBS show that in trees with high populations of A. tsugae , branches from the lower crown have slightly higher densities of A. tsugae than upper crown branches. In trees with low A. tsugae populations, the upper crown may have higher A. tsugae densities than the lower crown. North pointing branches also have higher densities of A. tsugae than branches pointing in other cardinal directions. Future sampling efforts for A. tsugae can take advantage of higher densities in certain portions of the crown to increase accuracy.     

沙地云杉构件动态与结构(英文)

研究了沙地云杉的构件种群。结果表明:(1)沙地云杉根系生长迅速;(2)针叶寿命随树龄而变化;(3)枝条数量与枝条年龄存在如下关系:B₄₀=2682.5e-0.6958A,B₂₀₀=5807.5e-1.3112A;(4)雄花/雌花随树龄迅速上升;(5)不同层次和方向上球果数量存在显著差异;(6)东方向的冠半径最长,西方向最短,枝条数量、长度,小枝数量随树龄而增加,小枝长度随树龄而缩短。结果表明,沙地云杉能够适应干旱的生态环境,其构件动态与结构是长期的自然选择与适应的结果。     

Architecture of Iberian canopy tree species in relation to wood density,shade tolerance and climate

Tree architecture has important consequences for tree performance as it determines resource capture, mechanical stability and dominance over competitors. We analyzed architectural relationships between stem and crown dimensions for 13 dominant Iberian canopy tree species belonging to the Pinaceae (six Pinus species) and Fagaceae (six Quercus species and Fagus sylvatica) and related these architectural traits to wood density, shade tolerance and climatic factors. Fagaceae had, compared with Pinaceae, denser wood, saplings with wider crowns and adults with larger maximal crown size but smaller maximal height. In combination, these traits enhance light acquisition and persistence in shaded environments; thus, contributing to their shade tolerance. Pinaceae species, in contrast, had low-density wood, allocate more resources to the formation of the central trunk rather than to branches and attained taller maximal heights, allowing them to grow rapidly in height and compete for light following disturbances; thus, contributing to their high light requirements. Wood density had a strong relationship with tree architecture, with dense-wooded species having smaller maximum height and wider crowns, probably because of cheaper expansion costs for producing biomechanically stable branches. Species from arid environments had shorter stems and shallower crowns for a given stem diameter, probably to reduce hydraulic path length and assure water transport. Wood density is an important correlate of variation in tree architecture between species and the two dominant families, with potentially large implications for their resource foraging strategies and successional dynamics.     

Variations in the foliar concentrations of macro and micro elements in a fast-growing tropical eucalypt

Summary The concentrations of N, P, K, Ca, Mg, Zn, Cu, Mn, Fe and B were measured in leaves of various ages in upper, mid and lower crown positions in Eucalyptus deglupta Blume during both wet and dry seasons. Based on coefficients of variation, the number of samples trees necessary for different levels of precision were calculated for each crown position.Least variation was found during the wet season for all elements except K. For all elements except Ca, fewest trees were needed when foliar material was collected from upper crown branches.The rate of leaf production in the upper crown was constant and it was possible to sample leaves of the same age by collecting from similar sampling positions; in contrast, that in the lower crowns was erratic and it was difficult to collect leaves of comparable age.The patterns of distribution and variation of foliar nutrients in the crown of E. deglupta are discussed. re]19750521     

Anomalous outbreaks of an invasive defoliator and native bark beetle facilitated by warm temperatures,changes in precipitation and interspecific interactions

Biotic disturbance agents such as insects can be highly responsive to climatic change and have widespread ecological and economic impacts on forests. Quantifying the responses of introduced and native insects to climate, including how dynamics of one agent may mediate those of another, is important for forecasting disturbance and associated impacts on forest structure and function. We investigated drivers of outbreaks by larch casebearer Coleophora laricella, an invasive defoliator, and eastern larch beetle Dendroctonus simplex, a native, tree‐killing bark beetle, on tamarack Larix laricina from 2000 to in Minnesota, USA. We evaluated the utility of temporal, spatial and climatic variables in predicting the presence/absence of outbreaks of each insect in cells of rasterized aerial survey data. The role of defoliation by larch casebearer in outbreaks of eastern larch beetle was also investigated. For both species, the most important predictors of outbreak occurrence were proximity of conspecific outbreaks in space and time. For larch casebearer, outbreak occurrence was positively associated with spring precipitation and warmer growing seasons. Outbreak occurrence of eastern larch beetle was positively associated with warmer and dryer years and was more likely in cells with prior defoliation by larch casebearer. Our results demonstrate that climate can drive large scale outbreaks of introduced and non‐native disturbance agents on a single host species, and that interactions at the tree level between such agents may scale up to manifest across large temporal and spatial scales.     

Crown structure and vertical foliage distribution in 4-year-old plantation-grown Eucalyptus pilularis and Eucalyptus cloeziana

Tree growth and form are both influenced by crown architecture and how it effects leaf distribution and light interception. This study examined the vertical distribution of foliage in 4-year-old plantation-grown Eucalyptus pilularis Sm. and E. cloeziana F. Muell. trees. Leaf area (LA) distribution was determined at two different sites using allometric approaches to determine LA in crown sections and for whole trees. Leaf area was distributed more towards the upper crowns when canopies had been closed for longer. Leaf area was also skewed more towards the upper crowns for Eucalyptus pilularis than E. cloeziana. These species differences were consistent with differences in vertical light availability gradients as determined by point quantum sensors. Leaf area of individual branches was highly correlated with branch cross-sectional area (CSA) and whole-tree LA was closely related to stem CSA. Branch-level allometric relationships were influenced by site and crown position. However, the general allometric equations between stem size and whole-tree leaf area could be applied across sites. Results from this study suggest that pruning of live branches in these species should follow species-specific guides for the timing and height of pruning to optimise the effects on stem growth and form.     

Relationships between tree size, crown shape, gender segregation and sex allocation in Pinus halepensis, a Mediterranean pine tree

Background and Aims

Sex allocation has been studied mainly in small herbaceous plants but much less in monoecious wind-pollinated trees. The aim of this study was to explore changes in gender segregation and sex allocation by Pinus halepensis, a Mediterranean lowland pine tree, within tree crowns and between trees differing in their size or crown shape.

Methods

The production of new male and female cones and sex allocation of biomass, nitrogen and phosphorus were studied. The relationship between branch location, its reproductive status and proxies of branch vigour was also studied.

Key Results

Small trees produced only female cones, but, as trees grew, they produced both male and female cones. Female cones were produced mainly in the upper part of the crown, and male cones in its middle and lower parts. Lateral branch density was correlated with the number of male but not female cones; lateral branches were more dense in large than in small trees and even denser in hemispherical trees. Apical branches grew faster, were thicker and their phosphorus concentration was higher than in lateral shoots. Nitrogen concentration was higher in cone-bearing apical branches than in apical vegetative branches and in lateral branches with or without cones. Allocation to male relative to female function increased with tree size as predicted by sex allocation theory.

Conclusions

The adaptive values of sex allocation and gender segregation patterns in P. halepensis, in relation to its unique life history, are demonstrated and discussed. Small trees produce only female cones that have a higher probability of being pollinated than the probability of male cones pollinating; the female-first strategy enhances population spread. Hemispherical old trees are loaded with serotinous cones that supply enough seeds for post-fire germination; thus, allocation to males is more beneficial than to females.     

臭冷杉生物量分配格局及异速生长模型

摘 要：臭冷杉是长白山阔叶红松林中重要针叶树种,采用整株收获法分析21株臭冷杉地上地下生物量分配格局。在枝条水平上采用样枝直径（BD）、样枝长度（BL）、样枝所在轮生枝位置（WP）建立活枝、针叶生物量异速生长模型,在植株水平上采用胸径（DBH）、树高（H）、年龄（Age）、树冠长度（CL）、树冠比率（CR）、南北向冠幅（CW1）、东西向冠幅（CW2）等变量建立树干木质、树皮、活枝、针叶、粗根及整株生物量模型。并利用逐步线性回归法获得不同器官生物量最优模型。结果表明：（1）活枝生物量主要集中在树冠中下层,针叶生物量集中在树冠中层。树冠中层和下层枝叶生物量无显著差异（p>0.05）;（2）21株臭冷杉地上生物量和地下生物量变动范围分别为1.026–506.047 kg/株和0.241–112.000 kg/株。粗根、活枝、针叶、树干木质、树皮及枯枝生物量占整株生物量的相对比例分别为18.68%、18.39%、12.02%、39.29%、8.70%和2.92%;（3）地上生物量与地下生物量呈显著线性相关（p<0.001）,拟合线性方程斜率为0.23;（4）枝条水平上,活枝生物量模型解释量超过95%,平均预测误差小于30%。与单变量（BD）活枝生物量模型相比,2变量（BD、BL）和3变量（BD、BL、WP）模型解释量分别提高1.2%和2.0%,平均预测误差分别下降6.26%和9.27%。针叶生物量相对较难预测,模型解释量仅为82.7%,平均预测误差接近50%,模型中增加BL 和WP变量并未提高针叶生物量的预测精度。活枝生物量与BD、BL、WP正相关,针叶生物量与BD正相关,与BL、WP负相关;（5）植株水平上,基于胸径的单变量模型可解释量大于90%,增加树高变量未能显著提高生物量模型的预测精度。年龄决定了臭冷杉的树干生物量,忽视年龄变量将会产生生物量预测误差。树冠特征是影响枝叶生物量预测精度的重要变量。综合考虑模型的可解释量及回归系数显著性可知,胸径是预测臭冷杉不同器官生物量的可靠变量。     

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.