Daily timings of sap production in sugar maple in Quebec, Canada.

Global warming is affecting plant phenology, with potential consequences on the dynamics of growth reactivation of sugar maple and the timings of maple syrup production. In this study, we assess the temperatures inducing the daily reactivation or cessation of sap production. We selected 19 sugarbushes across Quebec, Canada, using a tapping method associated with the tubing system, we recorded the daily timings of onset and ending of sap production during winter and spring 2018, and we associated the hourly temperatures at each site. Sap production occurred from mid-February to the end of April, starting on average between 10 and 11 AM, and ending from 6 to 8 PM. We observed a seasonal pattern in the onset and ending of sap production during spring, with the onset showing a greater change than the ending. Onset and ending of sap production occurred mostly under temperatures ranging between -2 and 2 °C. The production of sap in maple is closely related to circadian freeze-thaw cycles and occurs under nighttime and daytime temperatures fluctuating below and above 0 °C. The daily lengthening of the duration of sap production mirrors the changes in the timings of freeze and thaw events and can be explained by the physical properties of the water and the physiological processes occurring during growth reactivation. The ongoing warming will result in earlier and warmer springs, which may anticipate the cycles of freeze and thaw and advance sap production in sugar maple.

How plant allometry influences bud phenology and fruit yield in two Vaccinium species.

BACKGROUND AND AIMS: Understanding how plant allometry, plant architecture and phenology contribute to fruit production can identify those plant traits that maximize fruit yield. In this study, we compared these variables and fruit yield for two shrub species, Vaccinium angustifolium and Vaccinium myrtilloides, to test the hypothesis that phenology is linked to the plants' allometric traits, which are predictors of fruit production. METHODS: We measured leaf and flower phenology and the above-ground biomass of both Vaccinium species in a commercial wild lowbush blueberry field (Quebec, Canada) over a 2-year crop cycle; 1 year of pruning followed by 1 year of harvest. Leaf and flower phenology were measured, and the allometric traits of shoots and buds were monitored over the crop cycle. We hand-collected the fruits of each plant to determine fruit attributes and biomass. KEY RESULTS: During the harvesting year, the leafing and flowering of V. angustifolium occurred earlier than that of V. myrtilloides. This difference was related to the allometric characteristics of the buds due to differences in carbon partitioning by the plants during the pruning year. Through structural equation modelling, we identified that the earlier leafing in V. angustifolium was related to a lower leaf bud number, while earlier flowering was linked to a lower number of flowers per bud. Despite differences in reproductive allometric traits, vegetative biomass still determined reproductive biomass in a log-log scale model. CONCLUSIONS: Growing buds are competing sinks for non-structural carbohydrates. Their differences in both number and characteristics (e.g. number of flowers per bud) influence levels of fruit production and explain some of the phenological differences observed between the two Vaccinium species. For similar above-ground biomass, both Vaccinium species had similar reproductive outputs in terms of fruit biomass, despite differences in reproductive traits such as fruit size and number.

Exploring trees in three dimensions: VoxR, a novel voxel-based R package dedicated to analysing the complex arrangement of tree crowns.

Background: Interest in tree form assessments using the terrestrial laser scanner (TLS) has increased in recent years. Yet many existing methods are limited to small-sized trees, principally due to noise and occlusion phenomena. In this paper, a novel voxel-based program that is dedicated to the analyses of large tree structures is presented. The method is based on the assumption that architectural trait variations (i.e. branching angle, bifurcation ratio, biomass allocation, etc.) influence the way a tree explores space. This method uses the concept of space exploration that considers a voxel as a portion of space explored by the tree. Once the TLS scene is voxelized, the program provides tools that extract qualitative (geometrical) and quantitative (volumetric) metrics. These tools measure (1) voxel dispersion in three dimensions (3-D), (2) projections of the voxel cloud in 2-D and (3) multi-temporal changes within a single tree crown. Scope: To test algorithm capabilities of measuring larger tree architectural traits, two application studies were conducted using point clouds that were either generated by a tree growth simulation model, thereby allowing algorithm application in a perfectly controlled environment, or acquired in the field with a TLS device. The space exploration concept makes it possible to take advantage of the volumetric nature of voxels to compensate for occlusion. The hypothesis that large-sized voxels can be used to reduce occlusion in the original point cloud was tested, as well as the consequences of voxel size on quantification of tree volume and on precision of derived metrics. Conclusions: Results show that space exploration is well adapted to highlight architectural differences among trees. They also suggest that large-sized voxels are efficient for occlusion compensation at the expense of metrics precision in some cases. The best resolution to choose depending on the research objectives and quality of the TLS scan is discussed.

How fresh is maple syrup? Sugar maple trees mobilize carbon stored several years previously during early springtime sap-ascent.

While trees store substantial amounts of nonstructural carbon (NSC) for later use, storage regulation and mobilization of stored NSC in long-lived organisms like trees are still not well understood. At two different sites with sugar maple (Acer saccharum), we investigated ascending sap (sugar concentration, δ(13) C, Δ(14) C) as the mobilized component of stored stem NSC during early springtime. Using the bomb-spike radiocarbon approach we were able to estimate the average time elapsed since the mobilized carbon (C) was originally fixed from the atmosphere and to infer the turnover time of stem storage. Sites differed in concentration dynamics and overall δ(13) C, indicating different growing conditions. The absence of temporal trends for δ(13) C and Δ(14) C indicated sugar mobilization from a well-mixed pool with average Δ(14) C consistent with a mean turnover time (TT) of three to five years for this pool, with only minor differences between the sites. Sugar maple trees hence appear well buffered against single or even several years of negative plant C balance from environmental stress such as drought or repeated defoliation by insects. Manipulative investigations (e.g. starvation via girdling) combined with Δ(14) C measurements of this mobilized storage pool will provide further new insights into tree storage regulation and functioning.

Linking ice accretion and crown structure: towards a model of the effect of freezing rain on tree canopies.

BACKGROUND AND AIMS: Despite a longstanding interest in variation in tree species vulnerability to ice storm damage, quantitative analyses of the influence of crown structure on within-crown variation in ice accretion are rare. In particular, the effect of prior interception by higher branches on lower branch accumulation remains unstudied. The aim of this study was to test the hypothesis that intra-crown ice accretion can be predicted by a measure of the degree of sheltering by neighbouring branches. METHODS: Freezing rain was artificially applied to Acer platanoides L., and in situ branch-ice thickness was measured directly and from LiDAR point clouds. Two models of freezing rain interception were developed: 'IceCube', which uses point clouds to relate ice accretion to a voxel-based index (sheltering factor; SF) of the sheltering effect of branch elements above a measurement point; and 'IceTree', a simulation model for in silico evaluation of the interception pattern of freezing rain in virtual tree crowns. KEY RESULTS: Intra-crown radial ice accretion varied strongly, declining from the tips to the bases of branches and from the top to the base of the crown. SF for branches varied strongly within the crown, and differences among branches were consistent for a range of model parameters. Intra-crown variation in ice accretion on branches was related to SF (R(2) = 0·46), with in silico results from IceTree supporting empirical relationships from IceCube. CONCLUSIONS: Empirical results and simulations confirmed a key role for crown architecture in determining intra-crown patterns of ice accretion. As suspected, the concentration of freezing rain droplets is attenuated by passage through the upper crown, and thus higher branches accumulate more ice than lower branches. This is the first step in developing a model that can provide a quantitative basis for investigating intra-crown and inter-specific variation in freezing rain damage.

PypeTree: a tool for reconstructing tree perennial tissues from point clouds.

The reconstruction of trees from point clouds that were acquired with terrestrial LiDAR scanning (TLS) may become a significant breakthrough in the study and modelling of tree development. Here, we develop an efficient method and a tool based on extensive modifications to the skeletal extraction method that was first introduced by Verroust and Lazarus in 2000. PypeTree, a user-friendly and open-source visual modelling environment, incorporates a number of improvements into the original skeletal extraction technique, making it better adapted to tackle the challenge of tree perennial tissue reconstruction. Within PypeTree, we also introduce the idea of using semi-supervised adjustment tools to address methodological challenges that are associated with imperfect point cloud datasets and which further improve reconstruction accuracy. The performance of these automatic and semi-supervised approaches was tested with the help of synthetic models and subsequently validated on real trees. Accuracy of automatic reconstruction greatly varied in terms of axis detection because small (length < 3.5 cm) branches were difficult to detect. However, as small branches account for little in terms of total skeleton length, mean reconstruction error for cumulated skeleton length only reached 5.1% and 1.8% with automatic or semi-supervised reconstruction, respectively. In some cases, using the supervised tools, a perfect reconstruction of the perennial tissue could be achieved.

Assessing the potential of low-cost 3D cameras for the rapid measurement of plant woody structure.

Detailed 3D plant architectural data have numerous applications in plant science, but many existing approaches for 3D data collection are time-consuming and/or require costly equipment. Recently, there has been rapid growth in the availability of low-cost, 3D cameras and related open source software applications. 3D cameras may provide measurements of key components of plant architecture such as stem diameters and lengths, however, few tests of 3D cameras for the measurement of plant architecture have been conducted. Here, we measured Salix branch segments ranging from 2-13 mm in diameter with an Asus Xtion camera to quantify the limits and accuracy of branch diameter measurement with a 3D camera. By scanning at a variety of distances we also quantified the effect of scanning distance. In addition, we also test the sensitivity of the program KinFu for continuous 3D object scanning and modeling as well as other similar software to accurately record stem diameters and capture plant form (<3 m in height). Given its ability to accurately capture the diameter of branches >6 mm, Asus Xtion may provide a novel method for the collection of 3D data on the branching architecture of woody plants. Improvements in camera measurement accuracy and available software are likely to further improve the utility of 3D cameras for plant sciences in the future.

Biomass allocation and plant morphology explain the difference in shrub species abundance in a temperate forest.

In forested ecosystems, shrubs must succeed in persisting in low-light environments, while simultaneously having the ability to rapidly expand and occupy newly created canopy openings, yet little is known about the traits that make this possible. We hypothesize that shrub species that are abundant in the understory exhibit a specific set of functional traits that define their ability to persist during unfavorable periods and to rapidly exploit newly created habitats. We tested this by comparing field-measured functional traits such as biomass allocation, leaf display, crown morphology, and leaf traits, across individual size classes and two gap-forest environments of five shrub species. We observed significant differences in traits between species, size classes, and gap-forest environments. These differences were primarily related to biomass allocation traits, followed by leaf display, crown morphology, and leaf traits. Abundant shrubs like mountain maple (Acer spicatum) and hazelnut (Corylus cornuta) invested significantly more biomass in roots, had a larger total leaf area, and displayed leaves in a more efficient manner to intercept light. The high investment in root biomass can be interpreted as shrubs exploiting the persistence and colonization strategy through resprouting. Permanent sub-canopy status likely explains the importance of efficient leaf display, wherein abundant shrubs had a large leaf area with minimal support structures.

Reconstruction and analysis of a deciduous sapling using digital photographs or terrestrial-LiDAR technology.

BACKGROUND AND AIMS: To meet the increasing need for rapid and non-destructive extraction of canopy traits, two methods were used and compared with regard to their accuracy in estimating 2-D and 3-D parameters of a hybrid poplar sapling. METHODS: The first method consisted of the analysis of high definition photographs in Tree Analyser (TA) software (PIAF-INRA/Kasetsart University). TA allowed the extraction of individual traits using a space carving approach. The second method utilized 3-D point clouds acquired from terrestrial light detection and ranging (T-LiDAR) scans. T-LiDAR scans were performed on trees without leaves to reconstruct the lignified structure of the sapling. From this skeleton, foliage was added using simple modelling rules extrapolated from field measurements. Validation of the estimated dimension and the accuracy of reconstruction was then achieved by comparison with an empirical data set. KEY RESULTS: TA was found to be slightly less precise than T-LiDAR for estimating tree height, canopy height and mean canopy diameter, but for 2-D traits both methods were, however, fully satisfactory. TA tended to over-estimate total leaf area (error up to 50 %), but better estimates were obtained by reducing the size of the voxels used for calculations. In contrast, T-LiDAR estimated total leaf area with an error of <6 %. Finally, both methods led to an over-estimation of canopy volume. With respect to this trait, T-LiDAR (14·5 % deviation) greatly surpassed the accuracy of TA (up to 50 % deviation), even if the voxels used were reduced in size. CONCLUSIONS: Taking into account their magnitude of data acquisition and analysis and their accuracy in trait estimations, both methods showed contrasting potential future uses. Specifically, T-LiDAR is a particularly promising tool for investigating the development of large perennial plants, by itself or in association with plant modelling.

Contrasting Carbon Allocation Strategies of Ring-Porous and Diffuse-Porous Species Converge Toward Similar Growth Responses to Drought.

Extreme climatic events that are expected under global warming expose forest ecosystems to drought stress, which may affect the growth and productivity. We assessed intra-annual growth responses of trees to soil water content in species belonging to different functional groups of tree-ring porosity. We pose the hypothesis that species with contrasting carbon allocation strategies, which emerge from different relationships between wood traits and canopy architecture, display divergent growth responses to drought. We selected two diffuse-porous species (Acer saccharum and Betula alleghaniensis) and two ring-porous species (Quercus rubra and Fraxinus americana) from the mixed forest of Quebec (Canada). We measured anatomical wood traits and canopy architecture in eight individuals per species and assessed tree growth sensitivity to water balance during 2008-2017 using the standardized precipitation evapotranspiration index (SPEI). Stem elongation in diffuse-porous species mainly depended upon the total number of ramifications and hydraulic diameter of the tree-ring vessels. In ring-porous species, stem elongation mainly depended upon the productivity of the current year, i.e., number of vessels and basal area increment. Diffuse-porous and ring-porous species had similar responses to soil water balance. The effect of soil water balance on tree growth changed during the growing season. In April, decreasing soil temperature linked to wet conditions could explain the negative relationship between SPEI and tree growth. In late spring, greater water availability affected carbon partitioning, by promoting the formation of larger xylem vessels in both functional groups. Results suggest that timings and duration of drought events affect meristem growth and carbon allocation in both functional groups. Drought induces the formation of fewer xylem vessels in ring-porous species, and smaller xylem vessels in diffuse-porous species, the latter being also prone to a decline in stem elongation due to a reduced number of ramifications. Indeed, stem elongation of diffuse-porous species is influenced by environmental conditions of the previous year, which determine the total number of ramifications during the current year. Drought responses in different functional groups are thus characterized by different drivers, express contrasting levels of resistance or resilience, but finally result in an overall similar loss of productivity.

Minimum spring temperatures at the provenance origin drive leaf phenology in sugar maple populations.

Late frost can cause damage to trees, especially to the developing bud of broadleaf species in spring. Through long-term adaptation, plants adjust leaf phenology to achieve an optimal trade-off between growing season length and frost avoidance. In this study, we aim to assess ecotypic differentiation in leaf development of sugar maple populations planted in a common garden. A total of 272 sugar maple seedlings from 29 Canadian provenances were planted at the northern boundary of the natural range, and the phenological phases of bud and leaf development were monitored during spring 2019. The wide geographical area under evaluation showed a complex seasonal pattern of temperature, with spring warming occurring later in the north and close to the sea. Overall, leaf development lasted between 20 and 36 days, from the end of May to end of June. We observed different timings and rates of leaf development among provenances, demonstrating the occurrence of ecotypes in this species. Minimum April temperatures of the original sites were able to explain such differences, while maximum April temperatures were not significant. Seedlings from sites with colder minimum April temperatures completed leaf development earlier and faster. On average, leaf development diverged by up to 6 days among provenances, with minimum April temperatures ranging from -3 to 3 °C. Our results demonstrated that the avoidance of late spring frost is a driving force of leaf development in sugar maple populations. In the colder sites, the growing season is a limiting factor for tree growth. Thus, when thermal conditions become favorable in spring, an earlier growth reactivation and high metabolic activity ensure a fast leaf emission, which maximizes the period available for photosynthesis and growth. These patterns demonstrate the long-term phenological adaptation of sugar maple populations to local climatic conditions and suggest the importance of frost events for leaf development.

Complex Above- and Below-Ground Growth Responses of Two Urban Tree Species Following Root, Stem, and Foliage Damage-An Experimental Approach.

Urban trees are subjected to numerous biotic and mechanical damages, which can affect their growth rates and health. However, for most species, a systematic analysis of tree above- and below-ground growth reactions to a variety of damages is still lacking. Under a fully factorial experimental setup, using two common urban trees (Celtis occidentalis, Fraxinus pennsylvanica), we tested the effects of various degrees of frequently occurring damage as defoliation, root reduction, and stem injuries for a total of 18 treatments. We hypothesized that (i) an increasing amount of damage would proportionally negatively affect both root and stem growth; (ii) there would be a lag or lasting effect on growth; and (iii) both species would react similarly to the treatments. Contrary to our expectation, increasing levels of single or combined damage did not have an incremental effect on either stem or root growth. Although Celtis was significantly less vigorous than Fraxinus, it did not react strongly to damage treatments compared to the control. Interestingly, Celtis that experienced stem damage alone or in combination with other damages showed higher growth rates than the control. For Celtis, root injury was the treatment having the most impact, decreasing both root and stem growth consistently throughout the 5 years following treatments, whereas defoliation decreased growth only in the first 2 years. All damage treatments negatively affected stem and root growth of Fraxinus trees. Stem growth was affected the most by defoliation in the first year following the treatment, while root injury became the driving factor in subsequent years. For both species, stem injury showed the least influence on growth rates. The control and low-level damage treatments often affected growth rates in a similar way, suggesting that low-intensity stress triggers compensatory reactions stimulating photosynthetic rates and nutrient utilization. The slower-growing tree species, Celtis, showed a less negative reaction to all damage treatments compared to Fraxinus. This study illustrates that various types of above- and below-ground injuries do not have a simple additive effect on tree growth and that trees are capable of compensating for the loss of foliage, roots, or phloem to meet their metabolic demand.

The relative weight of ontogeny, topology and climate in the architectural development of three North American conifers.

Knowledge of plant architecture allows retrospective study of plant development, hence provides powerful tools, through modelling and simulation, to link this development with environmental constraints, and then predict its response to global change. The present study aims to determine some of the main endogenous and exogenous variables driving the architectural development of three North American conifers. We measured architectural traits retrospectively on the trunk, branches and twigs of whole tree crowns for each species: annual shoot length (ASL), needle length, branching patterns and reproduction organs (male and female). We fitted a partial least square (PLS) regression to explain each architectural trait with respect to topological, ontogenic and climatic variables. Results showed a significant weight of these three groups of variables for previous and current year, corresponding, respectively, to organogenesis and elongation. Topological and ontogenic variables had the greatest weight in models. Particularly, all architectural traits were strongly correlated with ASL. We highlighted a negative architectural response of two species to higher than average temperatures, whereas the third one took advantage of these higher temperatures to some degree. Tree architectural development weekly but significantly improved with higher precipitation. Our study underlines the strong weight of topology and ontogeny in tree growth patterns at twig and branch scales. The correlation between ASL and other tree architectural traits should be integrated into architectural development models. Climate variables are secondary in importance at the twig scale. However, interannual climate variations influence all axis categories and branching orders and therefore significantly impact crown development as a whole. This latter impact may increase with climate change, especially as climate affects architectural traits over at least 2 years, through organogenesis and elongation.

In vivo and in situ rhizosphere respiration in Acer saccharum and Betula alleghaniensis seedlings grown in contrasting light regimes.

A perfusive method combined with an open-system carbon dioxide measurement system was used to assess rhizosphere respiration of Acer saccharum Marsh. (sugar maple) and Betula alleghaniensis Britton (yellow birch) seedlings grown in 8-l pots filled with coarse sand. We compared in vivo and in situ rhizosphere respiration between species, among light regimes (40, 17 and 6% of full daylight) and at different times during the day. To compute specific rhizosphere respiration, temperature corrections were made with either species-specific coefficients (Q10) based on the observed change in respiration rate between 15 and 21 degrees C or an arbitrarily assigned Q10 of 2. Estimated, species-specific Q10 values were 3.0 and 3.4 for A. saccharum and B. alleghaniensis, respectively, and did not vary with light regime. Using either method of temperature correction, specific rhizosphere respiration did not differ either between A. saccharum and B. alleghaniensis, or among light regimes except in A. saccharum at 6% of full daylight. At this irradiance, seedlings were smaller than in the other light treatments, with a larger fine root fraction of total root dry mass, resulting in higher respiration rates. Specific rhizosphere respiration was significantly higher during the afternoon than at other times of day when temperature-corrected on the basis of an arbitrary Q10 of 2, suggesting the possibility of diurnal variation in a temperature-independent component of rhizosphere respiration.

Acclimation of Betula alleghaniensis Britton to moderate soil water deficit: small morphological changes make for important consequences in crown display.

In the context of the predicted increasing frequency of summer droughts in the northeastern deciduous forest of North America due to climate change, we investigated the acclimation capacity of yellow birch, an economically important native tree species, to soil water deficit. We carried out an integrated examination of allocation of biomass, leaf physiology, branching pattern and in situ 3D crown display. Potted seedlings were subjected to moderate soil water deficit for four consecutive months during their second growing season. Individuals under soil water deficit showed a 40% decrease in biomass accumulation but no change in the relative allocation of biomass to the different plant compartments. Net CO2 assimilation rates at leaf level decreased under water deficit (~15%) but could not alone explain the total reduction in growth, excluding the carbon starvation hypothesis. The observed reduction in net CO2 assimilation rates was related to a decrease in stomatal conductance and chlorophyll content. STARzen (in situ silhouette to total leaf area ratio; a proxy for light interception efficiency) decreased under soil water deficit due to shifts in biomass allocation within the branch compartment from long upper axes to short bottom axes. Despite the fact that the understanding of the processes involved in growth reduction and branching pattern alteration will need more attention in future research, we conclude that under water deficit yellow birch at young stages will: (i) experience a substantial loss of growth and biomass; and (ii) acclimate through architectural plasticity rather than through changes in the relative allocation of root biomass to enhance its water management.

Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability.

In a 4-year study, we investigated changes in leaf physiology, crown morphology and whole-tree biomass allocation in seedlings and saplings of shade-tolerant sugar maple (Acer saccharum Marsh.) and intermediate shade-tolerant yellow birch (Betula alleghaniensis Britt.) growing in natural understory light (0.5 to 35% of full sunlight) or in understory light reduced by 50% with shade nets to simulate the effect of gap closure. Leaf physiological parameters were mainly influenced by the light gradient, whereas crown morphological and whole-tree allocational parameters were mainly influenced by tree size. No single physiological, morphological or allocational trait was identified that could explain the difference in shade tolerance between the species. Yellow birch had higher growth rates, biomass allocation to branches and leaf physiological plasticity and lower crown morphological plasticity in unmodified understory light than sugar maple. Sugar maple did not display significant physiological plasticity, but showed variation with tree size in both crown morphology and whole-tree biomass allocation. When sugar maple was small, a greater proportion of whole-tree biomass was allocated to roots. However, physiological differences between the species decreased with decreasing light and most morphological and allocational differences tended to disappear with increasing tree size, suggesting that many species differences in shade-tolerance are expressed mainly during the seedling stage. Understory trees of both species survived for 4 years under shade nets, possibly because of higher plasticity when small and the use of stored reserves when taller.

In situ quantification of experimental ice accretion on tree crowns using terrestrial laser scanning.

In the eastern hardwood forests of North America ice storms are an important disturbance event. Ice storms strongly influence community dynamics as well as urban infrastructure via catastrophic branch failure; further, the severity and frequency of ice storms are likely to increase with climate change. However, despite a long-standing interest into the effects of freezing rain on forests, the process of ice accretion and thus ice loading on branches remains poorly understood. This is because a number of challenges have prevented in situ measurements of ice on branches, including: 1) accessing and measuring branches in tall canopies, 2) limitations to travel during and immediately after events, and 3) the unpredictability of ice storms. Here, utilizing a novel combination of outdoor experimental icing, manual measurements and terrestrial laser scanning (TLS), we perform the first in situ measurements of ice accretion on branches at differing heights in a tree crown and with increasing duration of exposure. We found that TLS can reproduce both branch and iced branch diameters with high fidelity, but some TLS instruments do not detect ice. Contrary to the expectations of ice accretion models, radial accretion varied sharply within tree crowns. Initially, radial ice accretion was similar throughout the crown, but after 6.5 hours of irrigation (second scanning) radial ice accretion was much greater on upper branches than on lower (∼factor of 3). The slope of the change in radial ice accretion along branches increased with duration of exposure and was significantly greater at the second scanning compared to the first. We conclude that outdoor icing experiments coupled with the use of TLS provide a robust basis for evaluation of models of ice accretion and breakage in tree crowns, facilitating estimation of the limiting breaking stress of branches by accurate measurements of ice loads.

Does shade improve light interception efficiency? A comparison among seedlings from shade-tolerant and -intolerant temperate deciduous tree species.

Here, we tested two hypotheses: shading increases light interception efficiency (LIE) of broadleaved tree seedlings, and shade-tolerant species exhibit larger LIEs than do shade-intolerant ones. The impact of seedling size was taken into account to detect potential size-independent effects on LIE. LIE was defined as the ratio of mean light intercepted by leaves to light intercepted by a horizontal surface of equal area. Seedlings from five species differing in shade tolerance (Acer saccharum, Betula alleghaniensis, A. pseudoplatanus, B. pendula, Fagus sylvatica) were grown under neutral shading nets providing 36, 16 and 4% of external irradiance. Seedlings (1- and 2-year-old) were three-dimensionally digitized, allowing calculation of LIE. Shading induced dramatic reduction in total leaf area, which was lowest in shade-tolerant species in all irradiance regimes. Irradiance reduced LIE through increasing leaf overlap with increasing leaf area. There was very little evidence of significant size-independent plasticity of LIE. No relationship was found between the known shade tolerance of species and LIE at equivalent size and irradiance.