Effects of growth rates, tree morphology and site conditions on longevity of Norway spruce in the northern Swiss Alps

Longevity of trees is known to be associated with growth rates, but also with tree morphology and spatial influences. However, very little quantitative information is available on the effects of these biotic and abiotic influences on maximum ages of trees. The objectives of this study were to investigate the trade-off between longevity and growth rates of Norway spruce (Picea abies) and to quantify the effects of tree morphology and abiotic site conditions on longevity of this species. Data were collected along different topographical and climatic gradients in a 20?×?25?km study area in the northern part of the Swiss Alps (Glarus). The ages of the more than 100 sampled dead Norway spruces ranged between 50 and 367?years. Longevity of these trees was negatively related to tree growth, i.e. slow-growing trees tended to grow older than fast-growing trees. Tree height was positively associated with longevity for both upper and lower storey trees. Longevity of lower storey trees was increased with large crown diameter, but decreased with long crown length. Upper storey trees growing at higher altitude tended to get older than at lower altitude. We conclude that the combined effects of growth rates, variability in site conditions and different traits of tree morphology determine tree longevity of Norway spruce in the Swiss Alps. Because longevity is tightly linked to mortality rates of tree populations, our study may improve our understanding of long-term processes of forest dynamics under current and future climate.     

Influence of crown class and shade tolerance on individual tree development during deciduous forest succession in Connecticut, USA

Sixty years of individual tree crown class records were used to elucidate the influence of crown class (dominant, codominant, intermediate, or suppressed), shade tolerance (intolerant, midtolerant, or tolerant), and their interactions on the probability of individual tree movement among crown classes. Trees were measured at 10 year intervals between 1927 and 1987, excluding 1947, on 364 nominal 0.01 ha plots. A total of 14 154 individual tree records were used in this analysis. The transition rates among crown classes for 30 year intervals (1927–1957 and 1957–1987) were examined for all combinations of antecedent crown classes and tolerance rankings. The distribution of ingrowth among tolerance rankings was also examined.

Mortality rates increased with decreasing crown class for all tolerance rankings, and mortality rates increased with decreasing tolerance. Compared with tolerant trees, midtolerant and intolerant trees had higher rates of ascension into dominant and codominant crown classes and exhibited higher persistence rates in the dominant crown class. These factors suggest that midtolerant and intolerant trees have an advantage over tolerant trees in the higher crown classes, In contrast, tolerant trees had the advantage in suppressed and intermediate crown classes, with lower mortality rates, higher persistence, and higher rates of crown class ascension than for midtolerant or intolerant trees. Crown class stratification was driven by the change in relative advantage of each tolerance ranking among the crown class.

Results of this study suggest that the canopy stratum (upper canopy, lower canopy, or mixed) affected by disturbance is as important as disturbance scale, intensity, and frequency in influencing the composition of the suppressed crown class. Because the suppressed crown class is in a constant state of high flux, with fewer than 30% of the suppressed trees remaining in the suppressed crown class for any 30 year period, a small change in the relative persistence or ascension rates among tolerance rankings, whether by a different disturbance or climate regime, could alter the proportion of tolerance rankings in the suppressed crown class and ultimately result in an alternative succession.     

Thinning and prescribed fire effects on overstory tree and snag structure in dry coniferous forests of the interior Pacific Northwest

Forest thinning and prescribed fires are practices used by managers to address concerns over ecosystem degradation and severe wildland fire potential in dry forests. There is some debate, however, about treatment effectiveness in meeting management objectives as well as their ecological consequences. The purpose of this study was to assess changes to forest stand structure following thinning and prescribed fire treatments, alone and combined, in the eastern Cascade Mountains of Washington State. Treatments were applied to 12 management units, with each treatment combination replicated three times (including untreated controls). Thinning modified forest structure by reducing overall tree density by >60% and canopy bulk density by 50%, and increased canopy base height by ∼4 m, thereby reducing susceptibility to crown fire. The prescribed fire treatment, conversely, did not appreciably reduce tree density or canopy fuel loading, but was effective at increasing the density of standing dead trees, particularly when combined with thinning (37 snags/ha increase). Prescribed fire effects were more pronounced when used in combination with thinning. Thinning was more reliable for altering stand structure, but spring burning was lower in intensity and coverage than desired and may have led to results that downplay the efficacy of fire to meet forest restoration goals.     

Stem sapwood permeability in relation to crown dominance and site quality in self-thinning fire-origin lodgepole pine stands

Stem sapwood hydraulic permeability, tree leaf area, sapwood basal area, earlywood to latewood ratio of annual rings, radial variation in hydraulic permeability and stem hydraulic capacity were examined in dominant (D), codominant (CD) and suppressed (SP) lodgepole pine (Pinus contorta Dougl. ex Loud.) trees growing on medium and poor sites. Hydraulic permeability on a sapwood area basis (ks) was lower in suppressed trees (0.71 x 10(-12) m2) compared to dominants (1.97 x 10(-12) m2) and codominants (1.79 x 10(-12) m2), and higher on medium than on poor sites. The leaf/sapwood area ratio (S) varied with crown dominance position (D > CD > SP) but not by site type. Leaf specific conductivity (kL) did not vary between crown classes or site types. The relationship between leaf area and stem hydraulic supply capacity (Q*) was strong, but differed among crown classes. Dominant trees and trees from the medium sites had a greater proportion of earlywood in outer rings of sapwood than suppressed trees. Sapwood permeability declined from the cambium to the sapwood-heartwood boundary in all samples, but the decline was more gradual in dominant trees compared to codominant and suppressed trees; differences in the radial variation in sapwood permeability may be related to differences in S. Sapwood permeability is positively related to crown dominance, whereas subdominant (CD and SP) trees have greater Q* in relation to leaf area, leading us to propose that this may give subdominant trees a survival advantage, slowing self-thinning.     

Pine mistletoe (Viscum album ssp. austriacum) contributes to Scots pine (Pinus sylvestris) mortality in the Rhone valley of Switzerland

In recent years unusual high mortality of Scots pine (Pinus sylvestris) has been observed in the Swiss Rhone Valley. The exact causes, however, are not known. At a 2‐ha monitoring plot, tree mortality and crown condition have been monitored since 1996. Between 1996 and 2004, 59% of the Scots pines died, most of them following the drought periods 1996–1998 and 2003–2004, while only 15% of the deciduous trees died. Crown transparency, needle discolouration, dead branch percentage, mistletoe (Viscum album ssp. austriacum) rating, Tomicus sp. shoot feeding, male flowering effect, tree stem diameter, crown shading and social tree class assessed in 1998 were used in a logistic regression model to predict tree mortality. Crown transparency, mistletoe rating and percentage of dead branches were found significant in the model and the probability of tree mortality increased with increasing rankings of these parameters. Needle discolouration could be used to substitute ‘dead branch percentage’ as predictor. While crown transparency increased with mistletoe rating, for trees in the same transparency class, trees with medium and heavy mistletoe infection were two to four times more likely to die than trees with no or only low mistletoe infection. For the surviving trees we found that trees with mistletoes showed a significantly higher increase in transparency in the year following a drought than trees without, while in a drought year the opposite was true. At the beginning of the observations no significant differences in transparency had been found between the trees with and without mistletoe. However, by the end of the observation period trees with mistletoe had significantly higher crown transparency. We conclude that mistletoe infection can be considered as both a predisposing factor for tree death, by increasing needle loss following drought and a contributing factor by increasing water stress during drought.     

Seasonal dynamics,especially autumnal retranslocation,of nitrogen and phosphorus in foliage of dominant and suppressed trees of beech,Fagus sylvatica

Seasonal changes in the N and P content of foliage in a young forest of Fagus sylvatica were measured. Leaves from branches of the upper and lower crown of dominant trees and from suppressed trees were compared. Nutrient retranslocation rates during senescence differed considerably between trees. This variation appeared not to be related to any differences in environmental factors or tree vigour, and was probably genetically induced. In dominant trees the most efficient retranslocation of N was recorded in the upper crown and probably resulted from higher leaf temperatures and a longer senescent period in the sun leaves than in the shade leaves. Phosphorus retranslocation efficiency was higher in suppressed trees than in dominant ones, but no such tendency was observed with N. The most obvious difference between leaves at different crown levels concerned the time at which P translocation began; an outflow of P from leaves in the lower crown began in June, while in the upper crown this outflow did not begin until September/October.     

Growth characteristics and reproductive output of dwarf mistletoe-infected Juniperus polycarpos in Iran

Dwarf mistletoes are parasitic flowering plants that infect conifers, resulting in substantial loss of growth and mortality. Recently, forest managers in Iran are contemplating whether infection of Juniperus polycarpos C. Koch forests by dwarf mistletoe, Arceuthobium oxycedri （DC.） M. Bieb, influences tree vigor and contributes to insuffieient natural regeneration. The present study aimed at assessing the severity of infection and its impact on growth and reproductive output of./., polycar- pos. Infected and uninfected trees （n =20 each） were selected for assess- ment of diameter, height, crown area, and crown volume as well as quantity and quality of cones and seeds. The severity of infection of trees was determined by Hawksworth＇s 6-class dwarf mistletoe rating （DMR） system. The DMR system revealed that 40% of the infected sample trees were lightly infected （DMR =1-2） and 60% were moderately infected （DMR =3--4）. Growth characteristics did not differ significantly （p 〉 0.05） between infected and uninfected trees. However, moderate infec- tion affected the reproductive output of./. polycarpos by significantly （p 〈0.05） reducing the mean number of cones per unit area of the crown, increasing the number of damaged seeds, and reducing seed size and seedgermination capacity. We conclude that reproductive output of J.. poly- carpos is more sensitive than growth characters to moderate infection by juniper dwarf mistletoe, and this might partly account for poor natural regeneration.     

Relationships between crown projected area and components of above-ground biomass in Norway spruce trees in even-aged stands: Empirical results and their interpretation

Sample tree material was reanalyzed in order to study the relationships between horizontal crown projected area and components of above-ground biomass in Norway spruce (Picea abies (L.) Karst.) trees growing in even-aged stands. The needle mass of dominant trees increased linearly with the increase in crown projected area, but in co-dominant and dominanted trees the increase in needle mass levelled off toward larger crown projected areas. The branch mass of dominant and co-dominant trees accumulated faster than linearly with increasing crown projected area, whereas in dominated trees an approximately linear relationship existed between these two variables. The increase in needle and branch mass per unit increase in crown projected area was highest in dominant trees and decreased to co-dominant and dominated trees, i.e. with tree position in the canopy. The stem mass accumulated obviously faster than linearly and similarly in all tree classes with the increase in crown projected area. The narrow crown shape indicated a high density of all components of above-ground biomass per unit of crown projected area.     

Rapid living crown reduction caused by Gremmeniella abietina affects foliar nutrient concentrations of Scots pine

The foliar chemistry of diseased and healthy trees was studied one growing season after severe reduction in living crown caused by Gremmeniella abietina in four young Scots pine (Pinus sylvestris) stands. Sample trees were chosen pairwise on the basis of the living crown length: a diseased tree with about 50% live crown reduction and a healthy tree in each pair. Fifteen elements were determined in the youngest healthy needles on the lateral top shoots of each sample tree. Diseased trees had higher foliar boron, manganese and sodium concentrations and lower magnesium, iron, zinc, copper, potassium, nitrogen and sulphur concentrations compared to the healthy trees. Foliar calcium, aluminium, phosphorus, carbon and hydrogen concentrations did not differ between the diseased and healthy trees, except for P and A1 in two of the stands when the stands were analysed separately. Significant correlations between the needle element concentrations and crown ratio (length of the living crown/tree height) were found especially for B (increasing B with decreasing crown ratio) and for Mg, Fe and Zn (decreasing concentrations with decreasing crown ratio). The effect of G. abietina-induced living crown reduction on tree nutrient status and the role of these mineral nutrients in the susceptibility are discussed.     

Changes in shoot allometry with increasing tree height in a tropical canopy species,Elateriospermum tapos

Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.     

Floral induction in Eucalyptus nitens

Eucalyptus nitens (Deane & Maiden) Maiden takes at least five years to initiate flower buds from seed and is an infrequent and light flowerer. Because this behavior constitutes a major impediment to breeding programs, we examined the mechanisms controlling floral induction in E. nitens, with the long-term aim of reducing generation time and increasing seed yield. Application of paclobutrazol reduced the concentration of endogenous gibberellic acid (GA) in apical tissue and enhanced the reproductive activity of grafted trees maintained outside over winter in Canberra, Australia. Grafts maintained in a warm greenhouse over winter did not produce flower buds, despite the paclobutrazol-induced reduction in GA concentration of the apical tissue. Exposing untreated grafts, which had been maintained over winter in a warm greenhouse, to low temperature the following spring reduced growth but did not induce flower bud production. Addition of GA(3) to paclobutrazol-treated grafts reduced the effect of paclobutrazol on reproductive activity.     

The effects of belowground resources on aboveground allometric growth in Bornean tree species

Tree height and crown allometries reflect adaptations for resource acquisition and structural stability, as well as plastic responses to a heterogeneous environment. While both light and soil resources limit growth and influence competitive responses in tropical forests, the effects of belowground resources on allometries are less understood, especially within the understory. To characterize outcomes of tree competition along an edaphic resource gradient, we quantified variation in height and crown allometries of six Bornean tree species from contrasting regeneration niches (light-demanding vs. shade-tolerant) on two soil habitats (clay-fine loam and sandy loam) within a 52-ha forest dynamics plot. Using empirically-fit allometric parameters and diameter growth rates from plot census data, we modeled tree height and crown area growth over the projected life span of each species. Based on resource competition theory, we hypothesized that tree species specializing on and populations of generalist species growing on the relatively moister, more fertile clay-fine loam soil habitat would have faster height and crown growth rates, compared to those on the sandy loam habitat, regardless of regeneration niche. Among soil specialists and within generalists of both genera, trees growing on clay-fine loam had taller stems and larger crowns at a given age and faster height and crown area growth rates at most sizes than trees on sandy loam. Differences in height and crown growth were driven by the faster diameter growth rates of trees on clay-fine loam, not by differences in height- and crown-diameter allometries, as trees on sandy loam were significantly taller at a given diameter, and differences in crown allometry were not consistent across soil habitats. Characterizing the height and crown growth responses of trees along resource gradients provides insight into the mechanisms that maintain diversity in tropical forests. Our results point to the importance of adaptive and plastic responses to both above and belowground resource availability in determining the allometric growth of trees and suggest that this diversity of responses may contribute tree species coexistence through competition-based trade-off mechanisms and variation in growth among individuals. Additionally, as the importance estimating natural carbon sequestration increases with the escalating effects of anthropogenic climate change, differences in tree growth and architecture across soil habitats also have implications for the approximation of forest carbon storage on heterogeneous tropical soils.     

长白落叶松人工林树冠形状的模拟

以长白山地区 2 6a生长白落叶松人工林为研究对象 ,采用枝解析的方法 ,测定了 2 5株林木 (直径 10 5～2 4 9cm)的树冠变量 ,并建立了预测树冠外侧形状的冠形模型。基于枝条着枝深度 (DINC)和林木变量所建立的树冠形状模型包括 :基径、枝长、着枝角度和弦长等预估模型。对于大小相同树木的主要枝条来讲 ,这些树冠变量是随着DINC的增加而增大 ;而林木的胸径 (DBH)和树高 (HT)变量很好地反映了不同大小树木的冠形变化。冠形预测模型预测效果良好 ,充分体现了树冠结构的变化趋势 :树冠形状在树冠的中上部呈抛物线体 ,而在树冠的下部则为近圆柱体。文中所建模型 ,可以合理地描述长白落叶松人工林的树冠形状及其变化规律。     

幼年赤桉胸径与冠幅、树高、材积的相关性分析

对试验地的1470株赤桉进行树高、胸径、冠幅的测量,并计算出样木的单株材积。把胸径分别和冠幅、树高及株材积进行相关性分析并且建立数学模型,用SPSS软件对所选模型进行曲线估计。结果表明：其中幂方程的 R2最大,F 值亦为最大,说明赤桉胸径与树高的幂关系显著,可确定赤桉胸径与树高的最优回归方程为H=1.804D0.673。胸径—冠幅,胸径—材积的最优模型分别为CW=0.674D0.561,V=0.0001614D2.341。分别对3组最优模型进行适应性检验,结果表明：材积的3个最优回归模型预测误差均在＆#177;3%以内,方程预测精度较高,可用于估算立木树高、冠幅、材积。     

Preformation in vegetative buds of pistachio (Pistacia vera): relationship to shoot morphology, crown structure and rootstock vigor

Effects of rootstock, shoot carbohydrate status, crop load and crown position on the number of preformed leaf primordia in the dormant terminal and lateral buds of mature and immature 'Kerman' pistachio (Pistacia vera L.) trees were investigated to determine if rootstock vigor is associated with greater shoot preformation. There was no significant variation in preformation related to the factors studied, suggesting strong genetic control of preformation in 'Kerman' pistachio. The growth differences observed among trees on different rootstocks were associated with greater stimulation of neoformed growth in trees on the more vigorous rootstocks. However, most annual extension growth in mature tree crowns was preformed, contrasting with the relatively high rate of neoformation found in young tree crowns. Large amounts of neoformed growth in young trees may allow the trees to become established quickly and secure resources, whereas predominantly preformed growth in mature trees may allow for continued crown expansion without outgrowing available resources. We hypothesized that the stimulation of neoformed growth by the more vigorous rootstocks is associated with greater resource uptake or transport, or both. Understanding the source of variation in shoot extension growth on different rootstocks has important implications for orchard management practices.     

Shoot growth and crown development: effect of crown position in three-dimensional simulations

Trees have been increasingly considered as modular organisms, with individual shoots forming autonomous units that respond semi-independently to their surrounding environment. However, there is evidence for fairly strict hormonal control of tree crown development. Studies on the hydraulic architecture of trees suggest a closer functional connection between shoots and crown development than is postulated by the theory of branch autonomy. We studied how shoot growth pattern influences growth and crown architecture in young Scots pine trees simulated by the LIGNUM model assuming that (a) the growth of a shoot mainly depends on its light climate and (b) the growth of a shoot is influenced by its position within the crown. We determined shoot position within the crown based on a recently developed vigor index. The vigor index compares the relative axis cross-sectional area from the base of the tree to each shoot and gives a value of 1 to the pathway of the greatest cross-sectional area. All other shoots attain values between 0 and 1 depending on their cross-sectional areas and the cross-sectional areas of the branches leading there from the main axis. The shoot light climate is characterized by annually intercepted photosynthetically active radiation. We compared the results from simulations (a) and (b) against an independent data set. The addition of a within-shoot position index (the vigor index) to our simulation (simulation b) resulted in a more realistic tree form than that obtained with simulation (a) alone. We discuss the functional significance of the results as well as the possibilities of using an index of shoot position in simulations of crown architecture.     

Comparative performances of new and existing indices of crown asymmetry: an evaluation using tall trees of Eucalyptus pilularis(Smith)

Over the past 50 years,crown asymmetry of forest trees has been evaluated through several indices constructed from the perspective of projected crown shape or displacement but often on an ad hoc basis to address specific objectives related to tree growth and competition,stand dynamics,stem form,crown structure and treefall risks.Although sharing some similarities,these indices are largely incoherent and non-comparable as they differ not only in the scale but also in the direction of their values in indicating the degree of crown asymmetry.As the first attempt at devising normative measures of crown asymmetry,we adopted a relative scale between 0 for perfect symmetry and 1 for extreme asymmetry.Five existing crown asymmetry indices(CAIs) were brought onto this relative scale after necessary modifications.Eight new CAIs were adapted from measures of circularity for digital images in computer graphics,indices of income inequality in economics,and a bilateral symmetry indicator in plant leaf morphology.The performances of the 13 CAIs were compared over different numbers of measured crown radii for 30 projected crowns of mature Eucalyptus pilularis trees through benchmarking statistics and rank order correlation analysis.For each CAI,the index value based on the full measurement of 36 evenly spaced radii of a projected crown was taken as the true value in the benchmarking process.The index(CAI_(13))adapted from the simple bilateral symmetry measure proved to be the least biased and most precise.Its performance was closely followed by that of three other CAIs.The minimum number of crown radii that is needed to provide at least an indicative measure of crown asymmetry is four.For more accurate and consistent measures,at least 6 or 8 crown radii are needed.The range of variability in crown morphology of the trees under investigation also needs to be taken into consideration.Although the CAIs are from projected crown radii,they can be readily extended to individual tree crown metrics that are now commonly extracted from LiDAR and other remotely sensed data.Adding a normative measure of crown asymmetry to individual tree crown metrics will facilitate the process of big data analytics and artificial intelligence in forestry wherever crown morphology is among the factors to be considered for decision making in forest management.     

Shoot production through cutting culture and micrografting from mature tree explants inAcacia tortilis (Forsk.) Hayne subsp.raddiana (Savi) Brenan

Shoot propagation from mature tree explants ofAcacia tortilis subsp.raddiana, a tropical tree legume, was studied. The effects of (i) the size of cuttings and (ii) a recut treatment of the donor trees, on axillary bud growth from multinodal horticultural cuttings are reported. Apices from seedlings, axillary buds from young lignified branches of mature trees, and axillary buds obtained through in vitro axillary branching from microcuttings isolated onto mature trees were all successfully micrografted on seedling rootstocks. A sterilization procedure for field explants is reported. Propagation via cuttings and via micrografting are discussed with regard to rejuvenation of mature donor trees.     

Adaptation of the LIGNUM model for simulations of growth and light response in Jack pine

LIGNUM is a whole tree model, developed for Pinus sylvestris in Finland, that combines tree metabolism with a realistic spatial distribution of morphological parts. We hypothesize that its general concepts, which include the pipe model, functional balance, yearly carbon budget, and a set of architectural growth rules, are applicable to all trees. Adaptation of the model to Pinus banksiana, a widespread species of economic importance in North America, is demonstrated.

Conversion of the model to Jack pine entailed finding new values for 16 physiological and morphological parameters, and three growth functions. Calibration of the LIGNUM Jack pine model for open grown trees up to 15 years of age was achieved by matching crown appearance and structural parameters (height, foliage biomass, aboveground biomass) with those of real trees. A sensitivity study indicated that uncertainty in the photosynthesis and respiration parameters will primarily cause changes to the net annual carbon gain, which can be corrected through calibration of the growth rate. The effect of a decrease in light level on height, biomass, total tree branch length, and productivity were simulated and compared with field data. Additional studies yielded insight into branch pruning, carbon allocation patterns, crown structure, and carbon stress. We discuss the value of the LIGNUM model as a tool for understanding tree growth and survival dynamics in natural and managed forests.