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1.  Responses of leaf structure and photosynthetic properties to intra-canopy light gradients: a common garden test with four broadleaf deciduous angiosperm and seven evergreen conifer tree species 
Oecologia  2012;170(1):11-24.
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.
PMCID: PMC3422461  PMID: 22349756
Plant functional types; Leaf plasticity; Shade acclimation; Evergreen leaves; Leaf mass-per-area
2.  Growth and posture control strategies in Fagus sylvatica and Acer pseudoplatanus saplings in response to canopy disturbance 
Annals of Botany  2011;107(8):1345-1353.
Background and Aims
Forest tree saplings that grow in the understorey undergo frequent changes in their light environment to which they must adapt to ensure their survival and growth. Crown architecture, which plays a critical role in light capture and mechanical stability, is a major component of sapling adaptation to canopy disturbance. Shade-adapted saplings typically have plagiotropic stems and branches. After canopy opening, they need to develop more erect shoots in order to exploit the new light conditions. The objective of this study was to test whether changes in sapling stem inclination occur after canopy opening, and to analyse the morphological changes associated with stem reorientation.
A 4-year canopy-opening field experiment with naturally regenerated Fagus sylvatica and Acer pseudoplatanus saplings was conducted. The appearance of new stem axes, stem basal diameter and inclination along the stem were recorded every year after canopy opening.
Key Results
Both species showed considerable stem reorientation resulting primarily from uprighting (more erect) shoot movements in Fagus, and from uprighting movements, shoot elongation and formation of relay shoots in Acer. In both species, the magnitude of shoot uprighting movements was primarily related to initial stem inclination. Both the basal part and the apical part of the stem contributed to uprighting movements. Stem movements did not appear to be limited by stem size or by stem growth.
Stem uprighting movements in shade-adapted Fagus and Acer saplings following canopy disturbance were considerable and rapid, suggesting that stem reorientation processes play a significant role in the growth strategy of the species.
PMCID: PMC3101137  PMID: 21444338
Advance regeneration; canopy gap; biomechanics; gravitropism; shade tolerance; Fagus sylvatica; Acer pseudoplatanus
3.  Exploring Ecological Significance of Tree Crown Plasticity through Three-dimensional Modelling 
Annals of Botany  2007;101(8):1221-1231.
Background and Aims
Morphogenetic plasticity may be as important as physiological plasticity in determining plant adaptability to changing environmental conditions. This study examines the importance of crown plasticity of trees in stands.
A three-dimensional forest simulator is used to explore the impact of crown shape plasticity on tree growth. Crown deformation is mediated through the local response to light and overall allometric constraints governing tree dimensions. By altering shape response parameters of Hevea brasiliensis the impact of increased or decreased plasticity is explored in a variety of competitive environments defined by various combinations of tree density and relative frequency of different strategies. The possible interactions between plasticity and growth rate and plasticity and below-ground competition are also explored.
Key Results
Crown plasticity confers competitive superiority in all cases studied. Interactions with other processes may downplay or enhance this competitive advantage.
Simulation results strongly suggest that crown plasticity does have a significant impact on tree performance in nature and that commonly observed crown shape deformation response of trees is of adaptive value.
PMCID: PMC2710270  PMID: 17720977
Crown plasticity; 3D simulation; individual-based model; competition
4.  Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings 
Annals of Botany  2009;104(6):1195-1205.
Backgrounds and Aims
Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.
The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.
Key Results
Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.
Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.
PMCID: PMC2766207  PMID: 19734164
Masting; shoot demography; short shoot; long shoot; temporal variation; spatial variation; leaf area; shoot length; resource allocation; Betula grossa
5.  Diversity of Flowering and Fruiting Phenology of Trees in a Tropical Deciduous Forest in India 
Annals of Botany  2006;97(2):265-276.
•Background and Aims In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species.
•Methods Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles.
•Key Results Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3–4 months) in dry-season and winter-flowering species, 6–9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized.
•Conclusions The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.
PMCID: PMC2803360  PMID: 16357055
Tropical tree phenology; flowering types; fruiting; asynchrony; leafless period; semi-evergreen species; summer flowering; summer leaf flushing
6.  Responses of Crown Development to Canopy Openings by Saplings of Eight Tropical Submontane Forest Tree Species in Indonesia: A Comparison with Cool-temperate Trees 
Annals of Botany  2006;97(4):559-569.
• 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.
PMCID: PMC2803653  PMID: 16399792
Cool-temperate trees; crown allometry; crown architecture; height growth; Indonesia; saplings; plasticity; tropical trees
7.  Do Interspecific Differences in Sapling Growth Traits Contribute to the Co-dominance of Acer saccharum and Fagus grandifolia? 
Annals of Botany  2007;101(1):103-109.
Background and Aims
Acer saccharum and Fagus grandifolia are among the most dominant late-successional tree species in North America. The influence of sapling growth responses to canopy gaps on the co-dominance of the two species in an old-growth forest in southern Quebec, Canada was examined. Two predictions were evaluated: (a) F. grandifolia is more shade tolerant than A. saccharum due to greater sapling leaf area and net production per sapling in closed-canopy conditions; and (b) the height growth rate of A. saccharum in canopy gaps is greater than that of F. grandifolia due to increased net production per sapling.
Sapling crown allometry, net production and height growth rates were compared between and within the two species in closed canopy vs. canopy gaps. Standardized major axis regression was used to analyse differences in crown allometry.
Key Results
F. grandifolia had greater crown projection, sapling leaf area and net production rate per sapling than A. saccharum in closed-canopy conditions. In response to canopy gaps, net production per sapling increased to the same degree in both species. The net production per sapling of F. grandifolia thus was much greater than that of A. saccharum in both canopy gap and closed-canopy conditions. The height growth rate of both species increased in canopy gaps, but the degree of increase was greater in F. grandifolia than in A. saccharum.
F. grandifolia regenerated more successfully than A. saccharum in both closed-canopy conditions and canopy gaps, which indicates that the co-dominance of the two species cannot be maintained simply by interspecific differences in shade tolerance and growth in gaps. Previous research showed that although Fagus and Acer shared dominance at this site, their relative dominance shifted with edaphic conditions. This suggests that the widespread co-dominance of the two species in eastern North American forests is maintained by the joint influence of canopy disturbance and species-specific responses to the heterogeneity of moisture and fertility regimes within forested landscapes.
PMCID: PMC2701832  PMID: 17942590
American beech; crown architecture; crown allometry; height growth rate; net production rate; saplings; sugar maple
8.  Timing of Canopy Closure Influences Carbon Translocation and Seed Production of an Understorey Herb, Trillium apetalon (Trilliaceae) 
Annals of Botany  2007;101(3):435-446.
Background and Aims
The light availability on a temperate, deciduous-forest floor varies greatly, reflecting the seasonal leaf dynamics of the canopy trees. The growth and/or reproductive activity of understorey plants should be influenced by the length of the high-irradiance period from snowmelt to canopy closure. The aim of the present study was to clarify how spring-blooming species regulate the translocation of photosynthetic products to current reproduction and storage organs during a growing season in accordance with the changing light conditions.
Growth pattern, net photosynthetic rate, seed production, and shoot and flower production in the next year of Trillium apetalon were compared between natural and experimentally shaded conditions. Furthermore, translocation of current photosynthetic products within plants was assessed by a labelled carbon-chase experiment.
Key Results
During the high-irradiance period, plants showed high photosynthetic ability, in which current products were initially used for shoot growth, then reserved in the rhizome. Carbon translocation to developing fruit occurred after canopy closure, but this was very small due to low photosynthetic rates under the darker conditions. The shading treatment in the early season advanced the time of carbon translocation to fruit, but reduced seed production in the current year and flower production of the next year.
Carbon translocation to the storage organ had priority over seed production under high-irradiance conditions. A shortened bright period due to early canopy closure effectively restricts carbon assimilation, which greatly reduces subsequent reproductive output owing to low photosynthetic products for fruit development and small carbon storage for future reproduction. As populations of this species are maintained by seedling recruitment, acceleration of canopy closure timing may influence the maintenance and dynamics of populations.
PMCID: PMC2701820  PMID: 18056055
13C labelling; canopy closure; carbon translocation; deciduous forest; light availability; photosynthesis; spring-bloomer; Trillium apetalon
9.  Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an Asian tropical dry karst forest 
Annals of Botany  2012;110(1):189-199.
Background and Aims
The co-occurring of evergreen and deciduous angiosperm trees in Asian tropical dry forests on karst substrates suggests the existence of different water-use strategies among species. In this study it is hypothesized that the co-occurring evergreen and deciduous trees differ in stem hydraulic traits and leaf water relationships, and there will be correlated evolution in drought tolerance between leaves and stems.
A comparison was made of stem hydraulic conductivity, vulnerability curves, wood anatomy, leaf life span, leaf pressure–volume characteristics and photosynthetic capacity of six evergreen and six deciduous tree species co-occurring in a tropical dry karst forest in south-west China. The correlated evolution of leaf and stem traits was examined using both traditional and phylogenetic independent contrasts correlations.
Key Results
It was found that the deciduous trees had higher stem hydraulic efficiency, greater hydraulically weighted vessel diameter (Dh) and higher mass-based photosynthetic rate (Am); while the evergreen species had greater xylem-cavitation resistance, lower leaf turgor-loss point water potential (π0) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, Am, and dry season π0. Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, Dh, as well as dry season π0. Both wood density and leaf density were closely correlated with leaf water-stress tolerance and Am.
The results reveal the clear distinctions in stem hydraulic traits and leaf water-stress tolerance between the co-occurring evergreen and deciduous angiosperm trees in an Asian dry karst forest. A novel pattern was demonstrated linking leaf longevity with stem hydraulic efficiency and leaf water-stress tolerance. The results show the correlated evolution in drought tolerance between stems and leaves.
PMCID: PMC3380589  PMID: 22585930
Tropical dry forest; karst; leaf habit; hydraulic conductivity; cavitation resistance; leaf water-stress tolerance; wood density; leaf density; phylogenetic independent contrasts
10.  Crown Plasticity and Competition for Canopy Space: A New Spatially Implicit Model Parameterized for 250 North American Tree Species 
PLoS ONE  2007;2(9):e870.
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.
Methodology/Principal Findings
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.
PMCID: PMC1964803  PMID: 17849000
11.  Trade-offs Between Seedling Growth and Survival in Deciduous Broadleaved Trees in a Temperate Forest 
Annals of Botany  2007;99(3):537-544.
Background and Aims
In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest.
The relationship between seedling survival and seedling relative growth rate in mass (RGRM) or height (RGRH) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes.
Key Results
Within each species, both RGRM and RGRH usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGRM and RGRH in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG.
The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.
PMCID: PMC2802958  PMID: 17242042
Coexistence; gap; gap size; habitat selection; habitat width; light; niche partitioning; relative growth rate; seed size; successional status
12.  Biomass expansion factor and root-to-shoot ratio for Pinus in Brazil 
The Biomass Expansion Factor (BEF) and the Root-to-Shoot Ratio (R) are variables used to quantify carbon stock in forests. They are often considered as constant or species/area specific values in most studies. This study aimed at showing tree size and age dependence upon BEF and R and proposed equations to improve forest biomass and carbon stock. Data from 70 sample Pinus spp. grown in southern Brazil trees in different diameter classes and ages were used to demonstrate the correlation between BEF and R, and forest inventory data, such as DBH, tree height and age. Total dry biomass, carbon stock and CO2 equivalent were simulated using the IPCC default values of BEF and R, corresponding average calculated from data used in this study, as well as the values estimated by regression equations. The mean values of BEF and R calculated in this study were 1.47 and 0.17, respectively. The relationship between BEF and R and the tree measurement variables were inversely related with negative exponential behavior. Simulations indicated that use of fixed values of BEF and R, either IPCC default or current average data, may lead to unreliable estimates of carbon stock inventories and CDM projects. It was concluded that accounting for the variations in BEF and R and using regression equations to relate them to DBH, tree height and age, is fundamental in obtaining reliable estimates of forest tree biomass, carbon sink and CO2 equivalent.
PMCID: PMC3196695  PMID: 21943243
allometry; carbon; regression; CDM; modeling
13.  Above-ground Biomass Investments and Light Interception of Tropical Forest Trees and Lianas Early in Succession 
Annals of Botany  2007;99(1):141-151.
Background and Aims
Crown structure and above-ground biomass investment was studied in relation to light interception of trees and lianas growing in a 6-month-old regenerating forest.
The vertical distribution of total above-ground biomass, height, diameter, stem density, leaf angles and crown depth were measured for individual plants of three short-lived pioneers (SLPs), four long-lived pioneers (LLPs) and three lianas. Daily light interception per individual Φd was calculated with a canopy model. The model was then used to estimate light interception per unit of leaf mass (Φleaf mass), total above-ground mass (Φmass) and crown structure efficiency (Ea, the ratio of absorbed vs. available light).
Key Results
The SLPs Trema and Ochroma intercepted higher amounts of light per unit leaf mass (Φleaf mass) because they had shallower crowns, resulting in higher crown use efficiency (Ea) than the other species. These SLPs (but not Cecropia) were also taller and intercepted more light per unit leaf area (Φarea). LLPs and lianas had considerably higher amounts of leaf mass and area per unit above-ground mass (LMR and LAR, respectively) and thus attained Φmass values similar to the SLPs (Φmass=Φarea×LAR). Lianas, which were mostly self-supporting, had light interception efficiencies similar to those of the trees.
These results show how, due to the trade-off between crown structure and biomass allocation, SLPs, and LLPs and lianas intercept similar amount of light per unit mass which may contribute to the ability of the latter two groups to persist.
PMCID: PMC2802976  PMID: 17210607
Bolivia; canopy model; crown structure; leaf mass ratio; lianas; light interception; pioneers; specific leaf area; tropical forest
14.  Elucidating the Population Dynamics of Japanese Knotweed Using Integral Projection Models 
PLoS ONE  2013;8(9):e75181.
Plant demographic studies coupled with population modeling are crucial components of invasive plant management because they inform managers when in a plant’s life cycle it is most susceptible to control efforts. Providing land managers with appropriate data can be especially challenging when there is limited data on potentially important transitions that occur belowground. For 2 years, we monitored 4 clonal Japanese knotweed (Polygonumcuspidatum) infestations for emergence, survival, shoot height until leaf senescence, dry shoot biomass after senescence, and rhizome connections for 424 shoots. We developed an integral projection model using both final autumn shoot height and shoot biomass as predictors of survival between years, growth from year to year, and number of rhizomes produced by a shoot (fecundity). Numbers of new shoots within an infestation (population growth rate λ) were projected to increase 13-233% in a year, with the greatest increase at the most frequently disturbed site. Elasticity analysis revealed population growth at 3 of the 4 sites was primarily due to ramet survival between years and to year-to-year growth in shoot height and shoot biomass. Population growth at the fourth site, the most disturbed, was due to the large production of new rhizomes and associated shoots. In contrast to previous studies, our excavation revealed that most of the shoots were not interconnected, suggesting rhizome production may be limited by the size or age of the plants, resource availability, disturbance frequency, or other factors. Future integration of plant population models with more data on belowground growth structures will clarify the critical stages in Japanese knotweed life cycle and support land managers in their management decisions.
PMCID: PMC3779190  PMID: 24073249
15.  Architecture of the Pruned Tree: Impact of Contrasted Pruning Procedures Over 2 Years on Shoot Demography and Spatial Distribution of Leaf Area in Apple (Malus domestica) 
Annals of Botany  2007;99(6):1055-1065.
Background and Aims
Demography and spatial distribution of shoots are rarely studied on pruned trees. The present 2-year study deals with the effect of pruning strategies on shoot demography and development, and consequences on the spatial distribution of leaf area in three architecturally contrasted — from type II to IV — apple cultivars: ‘Scarletspur Delicious’, ‘Golden Delicious’ and ‘Granny Smith’.
All trees were initially subjected during 5 years to Central Leader training with winter heading on all long shoots. For 2 years, half of the trees were further trained with Centrifugal training, where removal of flowering shoots — called extinction pruning — was carried out along the trunk and at the bottom of branches at flowering time. During these 2 years, shoot type (vegetative, inflorescence) and length, and the three-dimensional spatial distribution of all shoots were assessed with an electromagnetic digitizer.
Key Results
Shoot demography, frequency of transitions toward an inflorescence from either an inflorescence (bourse-over-bourse) or a vegetative shoot (trend toward flowering), and the number of bourse-shoots per bourse were strongly affected by cultivar, with little influence of tree manipulation. In contrast, the proportion of vegetative long shoots developing from previous year latent buds was significantly lower in Centrifugal-trained trees for the three cultivars. Canopy volume showed large variations between cultivars, but only that of ‘Granny Smith’ was affected by tree manipulation in the 2 years. Spatial distribution of shoots varied significantly according to cultivar and manipulation. In ‘Scarletspur Delicious’ and, to a lesser extent ‘Golden Delicious’, the distribution of vegetative and flowering shoots in the outer and the inner parts, respectively, was not affected by tree manipulation. In contrast, in ‘Granny Smith’, vegetative shoots were stimulated in the periphery of Central Leader trees, whereas flowering shoots were stimulated in the periphery of Centrifugal-trained trees.
In apple, the variability of responses to contrasted pruning strategies partly depends on the genetically determined growth and flowering habit of the cultivar.
PMCID: PMC3244342  PMID: 17416911
Malus domestica; apple; architecture; tree ideotype; shoot demography; shoot type; spatial pattern; centrifugal training; central leader; extinction pruning; reiteration
16.  Seedling Stage Strategies as a Means of Habitat Specialization in Herbaceous Plants 
PLoS ONE  2011;6(7):e23006.
The regeneration niche has been little investigated in studies of community assembly and plant distribution. We examined adaptive associations between seedling traits and habitat specialization. Two habitat contrasts were investigated across several evolutionary lineages of angiosperms: species specialized to forest vs. open habitats and to dry vs. wet habitats. We also tested whether effects of shade and drought vary independently or, alternatively, if shade may amplify effects on drought-stressed plants. Seedling response in terms of growth rate, height, slenderness, specific leaf area (SLA) and degree of elongation (longest internode; petiole or leaf-sheath depending on species' morphology) to light and watering treatments was assessed. We used a factorial design involving three light regimes and two watering frequencies. The open-shaded habitat contrast and the dry-wet habitat contrast were investigated using six and five pairs of congeneric species, respectively. The congeneric species pair design controlled for confounding effects of evolutionary history prior to divergence in habitat specialization. Seedling growth rate generally decreased with shade and reduced watering frequency. Plant height was generally largest at intermediate light. Specialization to shaded habitats was associated with a more conservative growth strategy, i.e. showing a more modest growth response to increasing light. Species from all habitats showed the highest relative elongation at intermediate light, except for the moist-habitat species, for which elongation increased with shade. Contrary to our expectations, species from dry habitats grew bigger than species from moist habitats in all treatments. SLA responded to the light treatment, but not to watering regime. The contrasting light and moisture conditions across habitats appear to not have selected for differences in SLA. We conclude that seedling phase strategies of resource allocation in temperate herbs contribute to their habitat specialization. Habitat-specific seedling strategies and trade-offs in response to resource availability and environmental conditions may be important to adaptive specialization.
PMCID: PMC3146528  PMID: 21829576
17.  Micropropagation and non-steroidal anti-inflammatory and anti-arthritic agent boswellic acid production in callus cultures of Boswellia serrata Roxb. 
Micropropagation through cotyledonary and leaf node and boswellic acid production in stem callus of a woody medicinal endangered tree species Boswellia serrata Roxb. is reported. The response for shoots, roots and callus formation were varied in cotyledonary and leafy nodal explants from in vitro germinated seeds, if inoculated on Murshige and Skoog’s (MS) medium fortified with cytokinins and auxins alone or together. A maximum of 8.0 ± 0.1 shoots/cotyledonary node explant and 6.9 ± 0.1 shoots/leafy node explants were produced in 91 and 88 % cultures respectively on medium with 2.5 μM 6-benzyladenine (BA) and 200 mg l−1 polyvinylpyrrolidone (PVP). Shoots treated with 2.5 μM IBA showed the highest average root number (4.5) and the highest percentage of rooting (89 %). Well rooted plantlets were acclimatized and 76.5 % of the plantlets showed survival upon transfer to field conditions. Randomly amplified polymorphic DNA (RAPD) analysis of the micropropagated plants compared with mother plant revealed true-to-type nature. The four major boswellic acid components in calluses raised from root, stem, cotyledon and leaf explants were analyzed using HPLC. The total content of four boswellic acid components was higher in stem callus obtained on MS with 15.0 μM IAA, 5.0 μM BA and 200 mg l−1 PVP. The protocol reported can be used for conservation and exploitation of in vitro production of medicinally important non-steroidal anti-inflammatory metabolites of B. serrata.
PMCID: PMC3550693  PMID: 24381442
Anti-arthritic; Anti-inflammatory; Boswellia serrata; Boswellic acid; Burseraceae; Callus; Indian olibanum; Micropropagation; Salai guggul; Steroids
18.  Cork Oak Vulnerability to Fire: The Role of Bark Harvesting, Tree Characteristics and Abiotic Factors 
PLoS ONE  2012;7(6):e39810.
Forest ecosystems where periodical tree bark harvesting is a major economic activity may be particularly vulnerable to disturbances such as fire, since debarking usually reduces tree vigour and protection against external agents. In this paper we asked how cork oak Quercus suber trees respond after wildfires and, in particular, how bark harvesting affects post-fire tree survival and resprouting. We gathered data from 22 wildfires (4585 trees) that occurred in three southern European countries (Portugal, Spain and France), covering a wide range of conditions characteristic of Q. suber ecosystems. Post-fire tree responses (tree mortality, stem mortality and crown resprouting) were examined in relation to management and ecological factors using generalized linear mixed-effects models. Results showed that bark thickness and bark harvesting are major factors affecting resistance of Q. suber to fire. Fire vulnerability was higher for trees with thin bark (young or recently debarked individuals) and decreased with increasing bark thickness until cork was 3–4 cm thick. This bark thickness corresponds to the moment when exploited trees are debarked again, meaning that exploited trees are vulnerable to fire during a longer period. Exploited trees were also more likely to be top-killed than unexploited trees, even for the same bark thickness. Additionally, vulnerability to fire increased with burn severity and with tree diameter, and was higher in trees burned in early summer or located in drier south-facing aspects. We provided tree response models useful to help estimating the impact of fire and to support management decisions. The results suggested that an appropriate management of surface fuels and changes in the bark harvesting regime (e.g. debarking coexisting trees in different years or increasing the harvesting cycle) would decrease vulnerability to fire and contribute to the conservation of cork oak ecosystems.
PMCID: PMC3386235  PMID: 22787521
19.  Size-dependent leaf area ratio in plant twigs: implication for leaf size optimization 
Annals of Botany  2009;105(1):71-77.
Background and Aims
Although many hypotheses have been proposed to explain variation in leaf size, the mechanism underlying the variation remains not fully understood. To help understand leaf size variation, the cost/benefit of twig size was analysed, since, according to Corner's rule, twig size is positively correlated with the size of appendages the twig bears.
An extensive survey of twig functional traits, including twig (current-year shoots including one stem and few leaves) and leaf size (individual leaf area and mass), was conducted for 234 species from four broadleaved forests. The scaling relationship between twig mass and leaf area was determined using standardized major axis regression and phylogenetic independent comparative analyses.
Key Results
Leaf area was found to scale positively and allometrically with both stem and twig mass (stem mass plus leaf mass) with slopes significantly smaller than 1·0, independent of life form and habitat type. Thus, the leaf area ratio (the ratio of total leaf area to stem or twig mass) decreases with increasing twig size. Moreover, the leaf area ratio correlated negatively with individual leaf mass. The results of phylogenetic independent comparativeanalyses were consistent with the correlations. Based on the above results, a simple model for twig size optimization was constructed, from which it is postulated that large leaf size–twig size may be favoured when leaf photosynthetic capacity is high and/or when leaf life span and/or stem longevity are long. The model's predictions are consistent with leaf size variation among habitats, in which leaf size tends to be small in poor habitats with a low primary productivity. The model also explains large variations in leaf size within habitats for which leaf longevity and stem longevity serve as important determinants.
The diminishing returns in the scaling of total leaf area with twig size can be explained in terms of a very simple model on twig size optimization.
PMCID: PMC2794065  PMID: 19864268
Allometry; leaf size; twig size; leaf area ratio; scaling relationship; broadleaved species
20.  Leaf traits, shoot growth and seed production in mature Fagus sylvatica trees after 8 years of CO2 enrichment 
Annals of Botany  2011;107(8):1405-1411.
Background and Aims
Masting, i.e. synchronous but highly variable interannual seed production, is a strong sink for carbon and nutrients. It may, therefore, compete with vegetative growth. It is currently unknown whether increased atmospheric CO2 concentrations will affect the carbon balance (or that of other nutrients) between reproduction and vegetative growth of forest species. In this study, reproduction and vegetative growth of shoots of mature beech (Fagus sylvatica) trees grown at ambient and elevated atmospheric CO2 concentrations were quantified. It was hypothesized that within a shoot, fruiting has a negative effect on vegetative growth, and that this effect is ameliorated at increased CO2 concentrations.
Reproduction and its competition with leaf and shoot production were examined during two masting events (in 2007 and 2009) in F. sylvatica trees that had been exposed to either ambient or elevated CO2 concentrations (530 µmol mol−1) for eight consecutive years, between 2000 and 2008.
Key Results
The number of leaves per shoot and the length of terminal shoots was smaller or shorter in the two masting years compared with the one non-masting year (2008) investigated, but they were unaffected by elevated CO2 concentrations. The dry mass of terminal shoots was approx. 2-fold lower in the masting year (2007) than in the non-masting year in trees growing at ambient CO2 concentrations, but this decline was not observed in trees exposed to elevated CO2 concentrations. In both the CO2 treatments, fruiting significantly decreased nitrogen concentration by 25 % in leaves and xylem tissue of 1- to 3-year-old branches in 2009.
Our findings indicate that there is competition for resources between reproduction and shoot growth. Elevated CO2 concentrations reduced this competition, indicating effects on the balance of resource allocation between reproduction and vegetative growth in shoots with rising atmospheric CO2 concentrations.
PMCID: PMC3101148  PMID: 21493641
Beech; carbon autonomy; CO2 enrichment; Fagus sylvatica; mast seeding; nitrogen; resource allocation; trade-off; vegetative growth
21.  Optimizing nitrogen economy under drought: increased leaf nitrogen is an acclimation to water stress in willow (Salix spp.) 
Annals of Botany  2011;108(7):1347-1353.
Background and Aims
The major objective was to identify plant traits functionally important for optimization of shoot growth and nitrogen (N) economy under drought. Although increased leaf N content (area basis) has been observed in dry environments and theory predicts increased leaf N to be an acclimation to drought, experimental evidence for the prediction is rare.
A pedigree of 200 full-sibling hybrid willows was pot-grown in a glasshouse in three replicate blocks and exposed to two water regimes for 3 weeks. Drought conditions were simulated as repeated periods of water shortage. The total leaf mass and area, leaf area efficiency (shoot growth per unit leaf area, EA), area-based leaf N content (NA), total leaf N pool (NL) and leaf N efficiency (shoot growth per unit leaf N, EN) were assessed.
Key Results
In the water-stress treatment, shoot biomass growth was N limited in the genotypes with low NL, but increasingly limited by other factors in the genotypes with greatest NL. The NA was increased by drought, and drought-induced shift in NA varied between genotypes (significant G × E). Judged from the EA–NA relationship, optimal NA was 16 % higher in the water-stress compared with the well-watered treatment. Biomass allocation to leaves and shoots varied between treatments, but the treatment response of the leaf : shoot ratio was similar across all genotypes.
It is concluded that N-uptake efficiency and leaf N efficiency are important traits to improve growth under drought. Increased leaf N content (area basis) is an acclimation to optimize N economy under drought. The leaf N content is an interesting trait for breeding of willow bioenergy crops in a climate change future. In contrast, leaf biomass allocation is a less interesting breeding target to improve yield under drought.
PMCID: PMC3197455  PMID: 21896572
Biomass allocation; biomass production; drought; leaf nitrogen; plant breeding; trait functionality
22.  Environmental and Ontogenetic Effects on Intraspecific Trait Variation of a Macrophyte Species across Five Ecological Scales 
PLoS ONE  2013;8(4):e62794.
Although functional trait variability is increasingly used in community ecology, the scale- and size-dependent aspects of trait variation are usually disregarded. Here we quantified the spatial structure of shoot height, branch length, root/shoot ratio and leaf number in a macrophyte species Potamogeton maackianus, and then disentangled the environmental and ontogenetic effects on these traits. Using a hierarchical nested design, we measured the four traits from 681 individuals across five ecological scales: lake, transect, depth stratus, quadrat and individual. A notable high trait variation (coefficient variation: 48–112%) was observed within species. These traits differed in the spatial structure, depending on environmental factors of different scales. Shoot height and branch length were most responsive to lake, transect and depth stratus scales, while root/shoot ratio and leaf number to quadrat and individual scales. The trait variations caused by environment are nearly three times higher than that caused by ontogeny, with ontogenetic variance ranging from 21% (leaf number) to 33% (branch length) of total variance. Remarkably, these traits showed non-negligible ontogenetic variation (0–60%) in each ecological scale, and significant shifts in allometric trajectories at lake and depth stratus scales. Our results highlight that environmental filtering processes can sort individuals within species with traits values adaptive to environmental changes and ontogenetic variation of functional traits was non-negligible across the five ecological scales.
PMCID: PMC3633840  PMID: 23626856
23.  Do individual-tree growth models correctly represent height:diameter ratios of Norway spruce and Scots pine? 
Forest Ecology and Management  2010;260(10):1735-1753.
Research highlights
▶ We examined four individual tree models in Europe: BWIN, Moses, Silva and Prognaus. ▶ We simulated growth of open-grown trees and on research plots for 15 or 30 years. ▶ Height:diameter ratios were correctly predicted by all four models. ▶ Height:diameter ratios were within the bounds of open grown trees and dense stands. ▶ They decreased with age and density; dominant trees had lower ratios than mean trees.
Height:diameter ratios are an important measure of stand stability. Because of the importance of height:diameter ratios for forest management, individual-tree growth models should correctly depict height:diameter ratios. In particular, (i) height:diameter ratios should not exceed that of very dense stands, (ii) height:diameter ratios should not fall below that of open-grown trees, (iii) height:diameter ratios should decrease with increasing spacing, (iv) height:diameter ratios for suppressed trees should be higher than ratios for dominant trees. We evaluated the prediction of height:diameter ratios by running four commonly used individual-tree growth models in central Europe: BWIN, Moses, Silva and Prognaus. They represent different subtypes of individual-tree growth models, namely models with and without an explicit growth potential and models that are either distance-dependent (spatial) or distance-independent (non-spatial). Note that none of these simulators predict height:diameter ratios directly. We began by building a generic simulator that contained the relevant equations for diameter increment, height increment, and crown size for each of the four simulators. The relevant measures of competition, site characteristics, and stand statistics were also coded. The advantage of this simulator was that it ensured that no additional constraint was being imposed on the growth equations, and that initial conditions were identical. We then simulated growth for a 15- and 30-year period for Austrian permanent research plots in Arnoldstein and in Litschau, which represent stands at different age-classes and densities. We also simulated growth of open-grown trees and compared the results to the literature. We found that the general pattern of height:diameter ratios was correctly predicted by all four individual-tree growth models, with height:diameter ratios above that of open-grown trees and below that of very dense stands. All models showed a decrease of height:diameter ratios with age and an increase with stand density. Also, the height:diameter ratios of dominant trees were always lower than that of mean trees. Although in some cases the observed and predicted height:diameter ratios matched well, there were cases where discrepancies between observed and predicted height:diameter ratios would be unacceptable for practical management predictions.
PMCID: PMC2987550  PMID: 21151352
Stand stability; Height:diameter ratio; Individual-tree growth model; Model evaluation; Open-grown trees
24.  Influence of Vegetation Structure on Lidar-derived Canopy Height and Fractional Cover in Forested Riparian Buffers During Leaf-Off and Leaf-On Conditions 
PLoS ONE  2013;8(1):e54776.
Estimates of canopy height (H) and fractional canopy cover (FC) derived from lidar data collected during leaf-on and leaf-off conditions are compared with field measurements from 80 forested riparian buffer plots. The purpose is to determine if existing lidar data flown in leaf-off conditions for applications such as terrain mapping can effectively estimate forested riparian buffer H and FC within a range of riparian vegetation types. Results illustrate that: 1) leaf-off and leaf-on lidar percentile estimates are similar to measured heights in all plots except those dominated by deciduous compound-leaved trees where lidar underestimates H during leaf off periods; 2) canopy height models (CHMs) underestimate H by a larger margin compared to percentile methods and are influenced by vegetation type (conifer needle, deciduous simple leaf or deciduous compound leaf) and canopy height variability, 3) lidar estimates of FC are within 10% of plot measurements during leaf-on periods, but are underestimated during leaf-off periods except in mixed and conifer plots; and 4) depth of laser pulse penetration lower in the canopy is more variable compared to top of the canopy penetration which may influence within canopy vegetation structure estimates. This study demonstrates that leaf-off lidar data can be used to estimate forested riparian buffer canopy height within diverse vegetation conditions and fractional canopy cover within mixed and conifer forests when leaf-on lidar data are not available.
PMCID: PMC3561319  PMID: 23382966
25.  Leaf-traits and growth allometry explain competition and differences in response to climatic change in a temperate forest landscape: a simulation study 
Annals of Botany  2011;108(5):885-894.
Background and Aims
The ability to simulate plant competition accurately is essential for plant functional type (PFT)-based models used in climate-change studies, yet gaps and uncertainties remain in our understanding of the details of the competition mechanisms and in ecosystem responses at a landscape level. This study examines secondary succession in a temperate deciduous forest in eastern China with the aim of determining if competition between tree types can be explained by differences in leaf ecophysiological traits and growth allometry, and whether ecophysiological traits and habitat spatial configurations among PFTs differentiate their responses to climate change.
A temperate deciduous broadleaved forest in eastern China was studied, containing two major vegetation types dominated by Quercus liaotungensis (OAK) and by birch/poplar (Betula platyphylla and Populus davidiana; BIP), respectively. The Terrestrial Ecosystem Simulator (TESim) suite of models was used to examine carbon and water dynamics using parameters measured at the site, and the model was evaluated against long-term data collected at the site.
Key Results
Simulations indicated that a higher assimilation rate for the BIP vegetation than OAK led to the former's dominance during early successional stages with relatively low competition. In middle/late succession with intensive competition for below-ground resources, BIP, with its lower drought tolerance/resistance and smaller allocation to leaves/roots, gave way to OAK. At landscape scale, predictions with increased temperature extrapolated from existing weather records resulted in increased average net primary productivity (NPP; +19 %), heterotrophic respiration (+23 %) and net ecosystem carbon balance (+17 %). The BIP vegetation in higher and cooler habitats showed 14 % greater sensitivity to increased temperature than the OAK at lower and warmer locations.
Drought tolerance/resistance and morphology-related allocation strategy (i.e. more allocation to leaves/roots) played key roles in the competition between the vegetation types. The overall site-average impacts of increased temperature on NPP and carbon stored in plants were found to be positive, despite negative effects of increased respiration and soil water stress, with such impacts being more significant for BIP located in higher and cooler habitats.
PMCID: PMC3177688  PMID: 21835816
Succession; watershed; ecophysiological trait; landscape carbon dynamics; temperate forests; competition; simulation; Quercus liaotungensis; Betula platyphylla; Populus davidiana

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