Canopy structure, which can be defined as the sum of the sizes, shapes and relative placements of the tree crowns in a forest stand, is central to all aspects of forest ecology. But there is no accepted method for deriving canopy structure from the sizes, species and biomechanical properties of the individual trees in a stand. Any such method must capture the fact that trees are highly plastic in their growth, forming tessellating crown shapes that fill all or most of the canopy space.
We introduce a new, simple and rapidly-implemented model–the Ideal Tree Distribution, ITD–with tree form (height allometry and crown shape), growth plasticity, and space-filling, at its core. The ITD predicts the canopy status (in or out of canopy), crown depth, and total and exposed crown area of the trees in a stand, given their species, sizes and potential crown shapes. We use maximum likelihood methods, in conjunction with data from over 100,000 trees taken from forests across the coterminous US, to estimate ITD model parameters for 250 North American tree species. With only two free parameters per species–one aggregate parameter to describe crown shape, and one parameter to set the so-called depth bias–the model captures between-species patterns in average canopy status, crown radius, and crown depth, and within-species means of these metrics vs stem diameter. The model also predicts much of the variation in these metrics for a tree of a given species and size, resulting solely from deterministic responses to variation in stand structure.
This new model, with parameters for US tree species, opens up new possibilities for understanding and modeling forest dynamics at local and regional scales, and may provide a new way to interpret remote sensing data of forest canopies, including LIDAR and aerial photography.
A better understanding of the relationship between stand structure and productivity is required for the development of: a) scalable models that can accurately predict growth and yield dynamics for the world's forests; and b) stand management regimes that maximize wood and/or timber yield, while maintaining structural and species diversity.
We develop a cohort-based canopy competition model (“CAIN”), parameterized with inventory data from Ontario, Canada, to examine the relationship between stand structure and productivity. Tree growth, mortality and recruitment are quantified as functions of diameter and asymmetric competition, using a competition index (CAIh) defined as the total projected area of tree crowns at a given tree's mid-crown height. Stand growth, mortality, and yield are simulated for inventoried stands, and also for hypothetical stands differing in total volume and tree size distribution.
For a given diameter, tree growth decreases as CAIh increases, whereas the probability of mortality increases. For a given CAIh, diameter growth exhibits a humped pattern with respect to diameter, whereas mortality exhibits a U-shaped pattern reflecting senescence of large trees. For a fixed size distribution, stand growth increases asymptotically with total density, whereas mortality increases monotonically. Thus, net productivity peaks at an intermediate volume of 100–150 m3/ha, and approaches zero at 250 m3/ha. However, for a fixed stand volume, mortality due to senescence decreases if the proportion of large trees decreases as overall density increases. This size-related reduction in mortality offsets the density-related increase in mortality, resulting in a 40% increase in yield.
Size-related variation in growth and mortality exerts a profound influence on the relationship between stand structure and productivity. Dense stands dominated by small trees yield more wood than stands dominated by fewer large trees, because the relative growth rate of small trees is higher, and because they are less likely to die.
Background and Aims
Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns.
Focus is given to relationships between tree height and top-shoot traits, which include the shoot's leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoot's stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan.
Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height.
In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.
Allometry; current-year leader shoot; hierarchical Bayesian model; pipe model; tree height; water stress
• Background and Aims The phenotypic plasticity of leaf life span in response to low resource conditions has a potentially large impact on the plant carbon budget, notably in evergreen species not subject to seasonal leaf shedding, but has rarely been well documented. This study evaluates the plasticity of leaf longevity, in terms of its quantitative importance to the plant carbon balance under limiting light.
• Methods Seedlings of four tropical tree species with contrasting light requirements (Alstonia scholaris, Hevea brasiliensis, Durio zibethinus and Lansium domesticum) were grown under three light regimes (full sunlight, 45 % sunlight and 12 % sunlight). Their leaf dynamics were monitored over 18 months.
• Results All species showed a considerable level of plasticity with regard to leaf life span: over the range of light levels explored, the ratio of the range to the mean value of life span varied from 29 %, for the least plastic species, to 84 %, for the most. The common trend was for leaf life span to increase with decreasing light intensity. The plasticity apparent in leaf life span was similar in magnitude to the plasticity observed in specific leaf area and photosynthetic rate, implying that it has a significant impact on carbon gain efficiency when plants acclimate to different light regimes. In all species, median survival time was negatively correlated with leaf photosynthetic capacity (or its proxy, the nitrogen content per unit area) and leaf emergence rate.
• Conclusions Longer leaf life spans under low light are likely to be a consequence of slower ageing as a result of a slower photosynthetic metabolism.
Alstonia scholaris; carbon balance; Durio zibethinus; Hevea brasiliensis; Lansium domesticum; leaf life span; light; plasticity
Tree mortality is a key process underlying forest dynamics and community assembly. Understanding how tree mortality is driven by simultaneous drivers is needed to evaluate potential effects of climate change on forest composition. Using repeat-measure information from c. 400,000 trees from the Spanish Forest Inventory, we quantified the relative importance of tree size, competition, climate and edaphic conditions on tree mortality of 11 species, and explored the combined effect of climate and competition. Tree mortality was affected by all of these multiple drivers, especially tree size and asymmetric competition, and strong interactions between climate and competition were found. All species showed L-shaped mortality patterns (i.e. showed decreasing mortality with tree size), but pines were more sensitive to asymmetric competition than broadleaved species. Among climatic variables, the negative effect of temperature on tree mortality was much larger than the effect of precipitation. Moreover, the effect of climate (mean annual temperature and annual precipitation) on tree mortality was aggravated at high competition levels for all species, but especially for broadleaved species. The significant interaction between climate and competition on tree mortality indicated that global change in Mediterranean regions, causing hotter and drier conditions and denser stands, could lead to profound effects on forest structure and composition. Therefore, to evaluate the potential effects of climatic change on tree mortality, forest structure must be considered, since two systems of similar composition but different structure could radically differ in their response to climatic conditions.
• Background and Aims Growth in trunk height in canopy openings is important for saplings. How saplings increase height growth in canopy openings may relate to crown architectural constraints. Responses of crown development to canopy openings in relation to trunk height growth were studied for saplings (0·2–2·5 m tall) of eight tropical submontane forest tree species in Indonesia. The results of this study were also compared with those of temperate trees in northern Japan.
• Methods The crown architecture differed among the eight tropical species, i.e. they had sparsely to highly developed branching structures. Crown allometry was compared among the eight species in each canopy condition (closed canopy or canopy openings), and between closed canopy and canopy openings within a species. A general linear regression model was used to analyse how each species increases height growth rate in canopy openings. Crown allometry and its plasticity were compared between tropical and temperate trees by a nested analysis of covariance.
• Key Results Tropical submontane trees had responses similar to cool-temperate trees, showing an increase in height in canopy openings, i.e. taller saplings of sparsely branched species increase height growth rates by increasing the sapling leaf area. Cool-temperate trees have a wider crown projection area and a smaller leaf area per crown projection area to avoid self-shading within a crown compared with tropical submontane trees. Plasticity of the crown projection area is greater in cool-temperate trees than in tropical submontane trees, probably because of the difference in leaf longevity.
• Conclusions This study concluded that interspecific variation in the responses of crown development to canopy openings in regard to increasing height related to the species' branching structure, and that different life-forms, such as evergreen and deciduous trees, had different crown allometry and plasticity.
Cool-temperate trees; crown allometry; crown architecture; height growth; Indonesia; saplings; plasticity; tropical trees
The Andes provide an extensive latitudinal and topographical framework for studying the factors that control the spatial patterns of forests (timberlines) and their species components expressed through the presence of tree growth forms (tree lines). Despite consistent overall similarities in landscape patterns, many processes must be unique, given the dramatic differences in species richness and biophysical constraints along the Andes. In all cases evaluated to date, morphological plasticity is a common trait of plant species that dominate at tree lines. In fact, many changes observed can be related to species-specific traits. Physiological limitations on tree growth form only explain species limits, while disturbances and cyclical climate fluctuations interact to affect many landscape patterns. Over long periods of time, tree lines provide unique habitats and perhaps opportunities for speciation. Understanding the spatial organization of tree-line dynamics is one viable research approach for evaluating the likely past fluxes and possible future changes.
Andes Mountains; biodiversity; timberline; tree line
Tree species differ from one another in, and display trade-offs among, a wide range of attributes, including canopy and understorey growth and mortality rates, fecundity, height and crown allometry, and crown transmissivity. But how does this variation affect the outcome of interspecific competition and hence community structure? We derive criteria for the outcome of competition among tree species competing for light, given their allometric and life-history parameters. These criteria are defined in terms of a new simple whole life-cycle measure of performance, which provides a simple way to organize and understand the many ways in which species differ. The general case, in which all parameters can differ between species, can produce coexistence, founder control or competitive exclusion: thus, competition for light need not be hierarchical as implied by R* theory. The special case in which species differ only in crown transmissivity produces neutral dynamics. The special case in which species differ in all parameters except crown transmissivity gives hierarchical competition, where the equivalent of R* is Zˆ*, the height at which trees enter the canopy in an equilibrium monoculture.
biomechanics; succession; community ecology; forest dynamics; pairwise invasibility; Sortie
Background and Aims
The dynamical system of plant growth GREENLAB was originally developed for individual plants, without explicitly taking into account interplant competition for light. Inspired by the competition models developed in the context of forest science for mono-specific stands, we propose to adapt the method of crown projection onto the x–y plane to GREENLAB, in order to study the effects of density on resource acquisition and on architectural development.
The empirical production equation of GREENLAB is extrapolated to stands by computing the exposed photosynthetic foliage area of each plant. The computation is based on the combination of Poisson models of leaf distribution for all the neighbouring plants whose crown projection surfaces overlap. To study the effects of density on architectural development, we link the proposed competition model to the model of interaction between functional growth and structural development introduced by Mathieu (2006, PhD Thesis, Ecole Centrale de Paris, France).
Key Results and Conclusions
The model is applied to mono-specific field crops and forest stands. For high-density crops at full cover, the model is shown to be equivalent to the classical equation of field crop production (
Howell and Musick, 1985, in Les besoins en eau des cultures; Paris: INRA Editions). However, our method is more accurate at the early stages of growth (before cover) or in the case of intermediate densities. It may potentially account for local effects, such as uneven spacing, variation in the time of plant emergence or variation in seed biomass. The application of the model to trees illustrates the expression of plant plasticity in response to competition for light. Density strongly impacts on tree architectural development through interactions with the source–sink balances during growth. The effects of density on tree height and radial growth that are commonly observed in real stands appear as emerging properties of the model.
Functional–structural plant models; GREENLAB; competition for light; Beer–Lambert Law; plant plasticity; dynamical system
As crowded populations of plants develop, the growth of some plants is accompanied by the death of others, a process called density-dependent mortality or 'self-thinning'. During the course of density-dependent mortality, the relationship between total population biomass (B) and surviving plant density (N) is allometric: B = aN(b). Essentially, increasing population biomass can be achieved only through decreasing population density. Variation in the allometric coefficient a among species has been recognized for many years and is important for management, assessment of productivity and carbon budgets, but the causes of this variation have not been elucidated. Individual-based models predict that size-dependent competition causes variation in the allometric coefficient. Using transgenic Arabidopsis with decreased plasticity, we provide experimental evidence that morphological plasticity of wild-type populations decreases the size asymmetry of competition for light and thereby decreases density-dependent mortality. This decrease in density-dependent mortality results in more biomass at a given density under size-symmetric compared with size-asymmetric competition.
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.
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.
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.
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.
Allocation; biomass; branch vigour; crown shape; gender segregation; nitrogen; Pinus halepensis; phosphorus; sex allocation; tree size
Subjective decisions of thematic and spatial resolutions in characterizing environmental heterogeneity may affect the characterizations of spatial pattern and the simulation of occurrence and rate of ecological processes, and in turn, model-based tree species distribution. Thus, this study quantified the importance of thematic and spatial resolutions, and their interaction in predictions of tree species distribution (quantified by species abundance). We investigated how model-predicted species abundances changed and whether tree species with different ecological traits (e.g., seed dispersal distance, competitive capacity) had different responses to varying thematic and spatial resolutions. We used the LANDIS forest landscape model to predict tree species distribution at the landscape scale and designed a series of scenarios with different thematic (different numbers of land types) and spatial resolutions combinations, and then statistically examined the differences of species abundance among these scenarios. Results showed that both thematic and spatial resolutions affected model-based predictions of species distribution, but thematic resolution had a greater effect. Species ecological traits affected the predictions. For species with moderate dispersal distance and relatively abundant seed sources, predicted abundance increased as thematic resolution increased. However, for species with long seeding distance or high shade tolerance, thematic resolution had an inverse effect on predicted abundance. When seed sources and dispersal distance were not limiting, the predicted species abundance increased with spatial resolution and vice versa. Results from this study may provide insights into the choice of thematic and spatial resolutions for model-based predictions of tree species distribution.
A recent study explored the adaptive significance of trunk inclination for trees growing on steep slopes. The authors used an optimality argument to predict how much a tree should respond to sidelight. Their calculations of the costs of leaning are flawed, because: (i) leaf mass has an allometric relationship with total volume, rather than being a fixed proportion of wood mass; (ii) the cost of support wood tissue is mainly a growth rate cost, not a maintenance respiration cost; (iii) small trees are sufficiently elastic to need very little support tissue, and thus they have a different risk structure; and (iv) most crown gaps are ephemeral rather than permanent, which also changes the risk equation. The argument I proposed in a previous study, addressing exactly the same question, is that canopy species are under selective pressure to maintain a strong central trunk that will reach the canopy and thus should not respond to sidelight. A reproductive value model is presented to illustrate this evolutionary question. Small, short-lived species or those from habitats with permanent openings (such as river margins) should be responsive. Both my previous data and the data of the other study support my model, but the model in the other study leaves many facts unexplained.
Trunk Inclination Leaf Biomass Shade-Tolerance
Models seldom consider the effect of leaf-level biochemical acclimation to temperature when scaling forest water use. Therefore, the dependence of transpiration on temperature acclimation was investigated at the within-crown scale in climatically contrasting genotypes of Acer rubrum L., cv. October Glory (OG) and Summer Red (SR). The effects of temperature acclimation on intracanopy gradients in transpiration over a range of realistic forest growth temperatures were also assessed by simulation. Physiological parameters were applied, with or without adjustment for temperature acclimation, to account for transpiration responses to growth temperature. Both types of parameterization were scaled up to stand transpiration (expressed per unit leaf area) with an individual tree model (MAESTRA) to assess how transpiration might be affected by spatial and temporal distributions of foliage properties. The MAESTRA model performed well, but its reproducibility was dependent on physiological parameters acclimated to daytime temperature. Concordance correlation coefficients between measured and predicted transpiration were higher (0.95 and 0.98 versus 0.87 and 0.96) when model parameters reflected acclimated growth temperature. In response to temperature increases, the southern genotype (SR) transpiration responded more than the northern (OG). Conditions of elevated long-term temperature acclimation further separate their transpiration differences. Results demonstrate the importance of accounting for leaf-level physiological adjustments that are sensitive to microclimate changes and the use of provenance-, ecotype-, and/or genotype-specific parameter sets, two components likely to improve the accuracy of site-level and ecosystem-level estimates of transpiration flux.
Intraspecific acclimation; MAESTRA; microclimate; modelling; red maple; temperature acclimation; transpiration
Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (Nmass) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (Narea). In evergreen conifers, LMA and Narea were less powerful predictors of area-based photosynthetic rate (Amax(area)) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, Amax(mass) was related to Nmass but not to LMA, whereas LMA became a significant codeterminant of Amax(mass) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.
Electronic supplementary material
The online version of this article (doi:10.1007/s00442-012-2279-y) contains supplementary material, which is available to authorized users.
Plant functional types; Leaf plasticity; Shade acclimation; Evergreen leaves; Leaf mass-per-area
Phenotypic plasticity is a key mechanism associated with the spread of exotic plants and previous studies have found that invasive species are generally more plastic than co-occurring species. Comparatively, the evolution of phenotypic plasticity in plant invasion has received less attention, and in particular, the genetic basis of plasticity is largely unexamined. Native from North America, Acer negundo L. is aggressively impacting the riparian forests of southern and eastern Europe thanks to higher plasticity relative to co-occurring native species. We therefore tested here whether invasive populations have evolved increased plasticity since introduction. The performance of 1152 seedlings from 8 native and 8 invasive populations was compared in response to nutrient availability. Irrespective of nutrients, invasive populations had higher growth and greater allocation to above-ground biomass relative to their native conspecifics. More importantly, invasive genotypes did not show increased plasticity in any of the 20 traits examined. This result suggests that the high magnitude of plasticity to nutrient variation of invasive seedlings might be pre-adapted in the native range. Invasiveness of A. negundo could be explained by higher mean values of traits due to genetic differentiation rather than by evolution of increased plasticity.
Background and Aims
Litter is a key factor in structuring plant populations, through positive or negative interactions. The litter layer forms a mechanical barrier that is often strongly selective against individuals lacking hypocotyle plasticity. Litter composition also interacts with plant growth by providing beneficial nutrients or, inversely, by allowing harmful allelopathic leaching. As conspicuous litter fall accumulation is often observed under deciduous forests, interactions between tree litter and understorey plant populations are worthy of study.
In a 1-year ex-situ experiment, the effects of tree litter on the growth of Anemone nemorosa, a small perennial forest geophyte, were investigated. Three ‘litter quantity’ treatments were defined, representative of forest floor litter (199, 356·5 and 514 g m−2), which were crossed with five ‘litter composition’ treatments (Quercus petraea, Fagus sylvatica, Carpinus betulus, Q. petraea + F. sylvatica and Q. petraea + C. betulus), plus a no-litter control. Path analysis was then used to investigate the pathways linking litter characteristics and components of adult plant growth.
As expected, the heavier the litter, the longer the petiole; rhizome growth, however, was not depreciated by the litter-induced petiole lengthening. Both rhizome mass increment and number of initiated buds marginally increased with the amount of litter. Rhizome mass increment was in fact determined primarily by leaf area and leaf life span, neither of which was unequivocally correlated with any litter characteristics. However, the presence of litter significantly increased leafing success: following a late frost event, control rhizomes growing in the absence of litter experienced higher leaf mortality before leaf unfolding.
The study questions the role of litter as a physical or chemical barrier to ground vegetation; to better understand this role, there is a need for ex-situ, longer-term experiments coupled with in-situ observations in the forest.
Anemone nemorosa; forest flora; geophyte; litter composition; litter permeability; path analysis; spring ephemeral; temperate forest
This paper documents the bizarre descending behavior from the tree crown to the ground of the larvae of the moth, Scopelodes contracta Walker (Lepidoptera: Limacodidae) and the interaction of the larva with predatory ants. S. contracta larvae infest leaves of many tree species in urban areas and orchards in Japan. Mature larvae and leaves without basal leaf parts were found under trees of four species infested with S. contracta larvae in Osaka, Japan. Individual larvae riding on leaves were observed falling from tree crowns to the ground. Many S. contracta cocoons were found in the soil below the trees two weeks after the observed parachuting. These observations indicate that S. contracta larvae parachuted to the ground where they spin their cocoons in the soil. When a larva that had just parachuted down was returned to an arboreal twig, the larva repeated the parachuting behavior. This parachuting behavior appears to be adaptive, because larvae can descend to the ground safely and with low energy cost. Worker ants of Tetramorium tsushimae Emery (Hymenoptera: Formicidae) and Pristomyrmex punctatus Mayr (Hymenoptera: Formicidae) occasionally attacked larvae on the ground before they had a chance to burrow in the soil.
descending behavior; larval movement; pupation site; slug caterpillar
▶ Determination of individual tree leaf area usually is only possible destructively. ▶ Surrogates which can be assessed non-destructively are investigated. ▶ From about 150 trees leaf area is estimated by 3P-branch sampling. ▶ These estimates are best correlated with crown surface area. ▶ Equations to determine individual tree leaf area non-destructively are presented.
Since individual tree leaf area is an important measure for productivity as well as for site occupancy, it is of high interest in many studies about forest growth. The exact determination of leaf area is nearly impossible. Thus, a common way to get information about leaf area is to use substitutes. These substitutes are often variables which are collected in a destructive way which is not feasible for long term studies. Therefore, this study aimed at testing the applicability of using substitutes for leaf area which could be collected in a non-destructive way, namely crown surface area and crown projection area. In 8 stands of Norway spruce (Picea abies L. Karst.), divided into three age classes and two thinning treatments, a total of 156 trees were felled in order to test the relationship between leaf area and crown surface area and crown projection area, respectively. Individual tree leaf area of the felled sample trees was estimated by 3P-branch sampling with an accuracy of ±10%. Crown projection area and crown surface area were compared with other, more commonly used, but destructive predictors of leaf area, namely sapwood area at different heights on the bole. Our investigations confirmed findings of several studies that sapwood area is the most precise measure for leaf area because of the high correlation between sapwood area and the leaf area. But behind sapwood area at crown base and sapwood area at three tenth of the tree height the predictive ability of crown surface area was ranked third and even better than that of sapwood area at breast height (R2 = 0.656 compared with 0.600). Within the stands leaf area is proportional to crown surface area. Using the pooled data of all stands a mixed model approach showed that additionally to crown surface area dominant height and diameter at breast height (dbh) improved the leaf area estimates. Thus, taking dominant height and dbh into account, crown surface area can be recommended for estimating the leaf area of individual trees. The resulting model was in line with many other findings on the leaf area and leaf mass relationships with crown size. From the additional influence of dominant height and dbh in the leaf area model we conclude that the used crown model could be improved by estimating the position of the maximum crown width and the crown width at the base of the crown depending on these two variables.
Norway spruce; Leaf area; Crown projection area; Crown surface area; Age; Thinning treatment
Debate continues in theoretical ecology over whether and why the scaling exponent of biomass–density (M–N) relationship varies along environmental gradients. By developing a novel geometric model with assumptions of allometric growth at the individual level and open canopy at the stand level, we propose that plant height–crown radius and canopy coverage–density relationships determine the above-ground M–N relationship in stressful environments. Results from field investigation along an aridity gradient (from eastern to western China) confirmed our model prediction and showed that the above-ground M–N scaling exponent increased with drought stress. Therefore, the ‘universal’ scaling exponents (−3/2 or −4/3) of the M–N relationship predicted by previous models may not hold for above-ground parts in stressful environments.
abiotic stress; allometric growth; aridity gradient; geometric model; self-thinning
The crown is the below ground portion of the stem of a grass which contains meristematic cells that give rise to new shoots and roots following winter. To better understand mechanisms of survival from freezing, a histological analysis was performed on rye, wheat, barley and oat plants that had been frozen, thawed and allowed to resume growth under controlled conditions. Extensive tissue disruption and abnormal cell structure was noticed in the center of the crown of all 4 species with relatively normal cells on the outside edge of the crown. A unique visual response was found in oat in the shape of a ring of cells that stained red with Safranin. A tetrazolium analysis indicated that tissues immediately inside this ring were dead and those outside were alive. Fluorescence microscopy revealed that the barrier fluoresced with excitation between 405 and 445 nm. Three dimensional reconstruction of a cross sectional series of images indicated that the red staining cells took on a somewhat spherical shape with regions of no staining where roots entered the crown. Characterizing changes in plants recovering from freezing will help determine the genetic basis for mechanisms involved in this important aspect of winter hardiness.
Background and Aims
Our knowledge about the influences of environmental factors on tree growth is principally based on the study of dominant trees. However, tree social status may influence intra-annual dynamics of growth, leading to differential responses to environmental conditions. The aim was to determine whether within-stand differences in stem diameters of trees belonging to different crown classes resulted from variations in the length of the growing period or in the rate of cell production.
Cambial activity was monitored weekly in 2006 for three crown classes in a 40-year-old silver-fir (Abies alba) plantation near Nancy (France). Timings, duration and rate of tracheid production were assessed from anatomical observations of the developing xylem.
Cambial activity started earlier, stopped later and lasted longer in dominant trees than in intermediate and suppressed ones. The onset of cambial activity was estimated to have taken 3 weeks to spread to 90 % of the trees in the stand, while the cessation needed 6 weeks. Cambial activity was more intense in dominant trees than in intermediate and suppressed ones. It was estimated that about 75 % of tree-ring width variability was attributable to the rate of cell production and only 25 % to its duration. Moreover, growth duration was correlated to tree height, while growth rate was better correlated to crown area.
These results show that, in a closed conifer forest, stem diameter variations resulted principally from differences in the rate of xylem cell production rather than in its duration. Tree size interacts with environmental factors to control the timings, duration and rate of cambial activity through functional processes involving source–sink relationships principally, but also hormonal controls.
Cambial activity; forest-stand structure; silver fir (Abies alba); tree-ring formation; tree-to-tree competition; social status; wood anatomy; xylem cell differentiation
Deformation behavior of a human dentin under compression including size and rate effects is studied. No difference between mechanical properties of crown and root dentin is found. It is mechanically isotropic high elastic and strong hard tissue, which demonstrates considerable plasticity and ability to suppress a crack growth. Mechanical properties of dentin depend on a shape of samples and a deformation rate.
Individual trees have been shown to exhibit strong relationships between DBH, height and volume. Often such studies are cited as justification for forest volume or standing biomass estimation through remote sensing. With resolution of common satellite remote sensing systems generally too low to resolve individuals, and a need for larger coverage, these systems rely on descriptive heights, which account for tree collections in forests. For remote sensing and allometric applications, this height is not entirely understood in terms of its location. Here, a forest growth model (SERA) analyzes forest canopy height relationships with forest wood volume. Maximum height, mean, H100, and Lorey's height are examined for variability under plant number density, resource and species. Our findings, shown to be allometrically consistent with empirical measurements for forested communities world-wide, are analyzed for implications to forest remote sensing techniques such as LiDAR and RADAR. Traditional forestry measures of maximum height, and to a lesser extent H100 and Lorey's, exhibit little consistent correlation with forest volume across modeled conditions. The implication is that using forest height to infer volume or biomass from remote sensing requires species and community behavioral information to infer accurate estimates using height alone. SERA predicts mean height to provide the most consistent relationship with volume of the height classifications studied and overall across forest variations. This prediction agrees with empirical data collected from conifer and angiosperm forests with plant densities ranging between 102–106 plants/hectare and heights 6–49 m. Height classifications investigated are potentially linked to radar scattering centers with implications for allometry. These findings may be used to advance forest biomass estimation accuracy through remote sensing. Furthermore, Lorey's height with its specific relationship to remote sensing physics is recommended as a more universal indicator of volume when using remote sensing than achieved using either maximum height or H100.
The genetic control of important adaptive traits, such as bud set, is still poorly understood in most forest trees species. Poplar is an ideal model tree to study bud set because of its indeterminate shoot growth. Thus, a full-sib family derived from an intraspecific cross of P. nigra with 162 clonally replicated progeny was used to assess the phenotypic plasticity and genetic variation of bud set in two sites of contrasting environmental conditions.
Six crucial phenological stages of bud set were scored. Night length appeared to be the most important signal triggering the onset of growth cessation. Nevertheless, the effect of other environmental factors, such as temperature, increased during the process. Moreover, a considerable role of genotype × environment (G × E) interaction was found in all phenological stages with the lowest temperature appearing to influence the sensitivity of the most plastic genotypes.
Descriptors of growth cessation and bud onset explained the largest part of phenotypic variation of the entire process. Quantitative trait loci (QTL) for these traits were detected. For the four selected traits (the onset of growth cessation (date2.5), the transition from shoot to bud (date1.5), the duration of bud formation (subproc1) and bud maturation (subproc2)) eight and sixteen QTL were mapped on the maternal and paternal map, respectively. The identified QTL, each one characterized by small or modest effect, highlighted the complex nature of traits involved in bud set process. Comparison between map location of QTL and P. trichocarpa genome sequence allowed the identification of 13 gene models, 67 bud set-related expressional and six functional candidate genes (CGs). These CGs are functionally related to relevant biological processes, environmental sensing, signaling, and cell growth and development. Some strong QTL had no obvious CGs, and hold great promise to identify unknown genes that affect bud set.
This study provides a better understanding of the physiological and genetic dissection of bud set in poplar. The putative QTL identified will be tested for associations in P. nigra natural populations. The identified QTL and CGs will also serve as useful targets for poplar breeding.