Solar equivalent leaf area: an efficient biometrical parameter of individual leaves, trees and stands

The solar equivalent leaf area (A(s)), a simply and easily determined biometrical parameter of leaves, trees and stands, was derived theoretically. The parameter is defined as projected leaf area weighted for the time integral of irradiance at a given location in the canopy relative to that of fully irradiated leaves at the top of the canopy. The efficiency of A(s) as a basis for estimating stand-area transpiration of a mature oak (Quercus robur L.) forest from measurements of transpiration by individual trees was compared with that of other stand and tree characteristics. Stand transpiration estimates based on A(s) were more precise and less prone to systematic error than estimates based on basal area, timber volume, projected tree crown area, projected leaf area, or leaf dry mass. Solar equivalent leaf area reflects both the amount and the physiological properties of leaves and can be used as a measure of tree size and functional capacity. It can be calculated from ordinary forest inventory data on trees and stands, adjusted according to simple phyllometric data. It appears to have wide application in ecological and forestry studies for relating the physiological characteristics of individual leaves to those of entire trees or stands.     

日本栗叶面积与叶片生物量计算方法的初步研究

以6年生日本栗品种＂辽栗10号＂与＂大峰＂为试材,对其叶长、叶宽、单叶面积、单叶干重、冠幅、冠高、单株叶片生物量进行了测定。研究不同品种间叶片长宽比值和比叶面积的差异性,叶片长、叶宽和叶片长宽乘积与单叶面积的相关性以及冠幅、冠高与单株叶片生物量的相关性。结果表明：不同品种间叶片长宽比、比叶面积均无显著差异;对日本栗叶长、叶宽、叶片长宽乘积与单叶面积进行一元回归方程拟合,决定系数与修正决定系数均达极显著水平（P〈0.0001）,其中以叶片长宽乘积与单叶面积的一元二次回归方程回归效果最好,决定系数达0.987 9,修正决定系数达0.987 7;单株叶片生物量与冠幅面积和冠高的二元线性回归关系达极显著水平（P〈0.0001）,决定系数达0.997 2,修正决定系数达0.996 4。在生产上,可以通过测量冠幅面积、冠高计算出单株叶片生物量和叶面积指数,该方法操作简单,具有较高应用价值。     

Allometric estimation of above-ground biomass and leaf area in managed Grevillea robusta agroforestry systems

Non-destructive methods for determining the biomass and leaf area of individual trees throughout their growing cycle are an essential tool in agroforestry research, but must be capable of providing reliable estimates despite the influence that management strategies such as pruning may have on tree form. In the present study, allometric methods involving measurements of the diameter of all branches provided reliable estimates of canopy leaf area and biomass for grevillea trees (Grevillea robusta A. Cunn.; Proteaceae) grown as poles, but proved unsuitable for routine measurements because of their time-consuming nature. An alternative, less laborious method based on measurements of trunk cross-sectional area immediately below the first branch of the canopy provided satisfactory allometric estimates of leaf area and canopy biomass. Trunk biomass was determined from measurements of tree height and diameter at breast height using established methodology based on the assumption that trunk volume may be calculated using a quadratic paraboloid model; biomass was determined as the product of trunk volume and the specific gravity of the wood. The theoretical basis, development and validation of allometric methods for estimating tree growth are discussed and their wider applicability to other agroforestry systems is assessed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon

Relationships between tree height and crown dimensions and trunk diameter were determined for shade-tolerant species of old-growth forests of western Oregon. The study included both understory and overstory species, deciduous and evergreen angiosperms and evergreen conifers. A comparison of adult understory species with sapling overstory species of similar height showed greater crown width and trunk diameter in the former, whether the comparison is made among conifers or deciduous trees. Conifer saplings had wider crowns than deciduous saplings, but the crown widths of the two groups converged with increase in tree height. Conifer saplings had thicker trunks than deciduous saplings of similar crown width, possibly because of selection for resistance to stem bending under snow loads. The results suggest that understory species have morphologies that increase light interception and persistence in the understory, whereas overstory species allocate their biomass for efficient height growth, thereby attaining the high-light environment of the canopy. The greater crown widths and the additional strength requirements imposed by snow loads on conifer saplings result in less height growth per biomass increment in conifer saplings than in deciduous saplings. However, the convergence in crown width of the two groups at heights greater than 20 m, and the proportionately smaller effect of snow loads on large trees, may result in older conifers equalling or surpassing deciduous trees in biomass allocation to height growth.     

Modelling biomass and leaf area index in a sub-arctic Scandinavian mountain area

Estimates of biomass and leaf area index (LAI) are important variables in ecological and climate models. However, very little is known about the biomass and LAI of the vegetation in the Scandinavian mountain area. In this study, extensive field data consisting of diameter at breast height for 13?000 trees and height for 550 trees were collected. Furthermore, biomass and leaf area (LA) measurements for 46 mountain birch trees [Betula pubescens ssp. czerepanovii (Orlowa) Hämet-Ahti] and biomass and LA measurements for shrubs (e.g. Salix spp., Betula nana) at 36 sample plots were carried out. Multiplicative linear models for trees were fitted to tree biomass and LA measurements using basal area at breast height, height, crown diameter and diameter at stump height as explanatory variables. Additive linear models were fitted to shrub biomass and LAI measurements using coverage of shrubs, topographic variables and soil type as explanatory variables. The functions were then used to predict the biomass and LAI for trees and shrubs for the entire test area, which covers an area of 84 km² and is located at latitude 68° N. The mean total biomass estimates were 27?493 kg ha^?1 for the forest and 7650 kg ha^?1 for snow-protected heath and meadow vegetation. The LAIs were 2.06 and 0.52, respectively. For monitoring biomass and LAI in the Scandinavian mountain area, the functions could also be applied to data from traditional field-based inventories and the estimates might further be improved by combining the estimates from the test area with auxiliary information such as remote sensing images.     

基于管道模型理论的长白落叶松叶生物量与径向生长关系研究

目的 为了解长白落叶松单木叶生物量与径向生长之间的关系。 方法 以小兴安岭地区长白落叶松为研究对象,结合管道模型理论,研究单木个体水平和单木内不同方位区间水平上的叶生物量与径向生长的关系,分析树冠分布与径向生长之间的相关性。 结果 叶生物量与胸高处和枝下高处树干直径、断面积、边材面积,近1、2、3、5年断面积生长量均呈显著正相关,使用胸高断面积作为预测因子的叶生物量模型拟合效果最好。单木株内不同方位区域的叶生物量与对应的胸径处和枝下高处的树干半径、断面积,近1、2、3、5年断面积生长量均呈显著正相关。 结论 估测单株叶生物量时,使用胸高断面积作为预测因子的预测精度最高。株内不同方位叶生物量与对应方位的树干半径、断面积及断面积生长量均呈显著正相关,树冠偏冠与髓心偏心具有一定的相关性。     

Applicability of non-destructive substitutes for leaf area in different stands of Norway spruce (Picea abies L. Karst.) focusing on traditional forest crown measures

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.     

Canopy fuel characteristics in relation to crown fire potential in pine stands: analysis, modelling and classification

Crown fire occurrence and subsequent crown fire behaviour are strongly dependent on canopy fuel characteristics, especially canopy fuel load (CFL), canopy bulk density (CBD) and canopy base height (CBH). Therefore, quantification of such variables is required for the appropriate selection of silvicultural treatments aimed at reducing susceptibility to crown fire. Data from the IV Spanish National Forest Inventory and individual tree biomass dry weight equations were used to estimate the canopy fuel characteristics of four representative types of pine stands in north-western Spain. Probability of crown fire initiation and crown fire rate of spread were simulated by using the mean surface fuel load observed for each type of pine in this area and assuming different burning conditions. The results indicate that a 22.13 % of the sample plots analysed showed a rather high potential for active crown fire spread under moderate burning conditions, and this value increases to 69.27 % under extreme burning conditions. Equations relating the canopy fuel characteristics to common stand variables (stand density, basal area and dominant height) were fitted simultaneously for each pine, and weighting factors for heteroscedasticity were included. The models explained more than 93.90, 74.70 and 69.42 % of the observed variability in CFL, CBD and CBH, respectively. Basal area was the most important variable for estimating CFL and CBD while dominant height explained most of the observed variability in CBH. The use of the fitted equations together with existing dynamic growth models and fire management decision support systems will enable assessment of the crown fire potential associated with different silvicultural alternatives used in these types of pine stands.     

Crown structure and leaf area index development in thinned and unthinned Eucalyptus nitens plantations

The crown structure of Eucalyptus nitens (Deane & Maiden) Maiden 6 years after thinning, and the development of stand leaf area index both immediately and 6 years after thinning, were investigated. Thinning did not alter branch angle, branching density or the relationship between branch size and branch leaf area. However, larger branches were found in the lower crown of thinned trees and the increase in leaf area as a result of thinning occurred on the northern aspect of the crown. The vertical distribution of leaf area in unthinned trees was skewed toward the top of the crown and correlated with live crown ratio. The vertical distribution of leaf area in thinned trees tended to be less skewed and was unrelated to tree size or dominance. Leaf area index, as estimated from light interception measurements, increased at a constant rate soon after thinning regardless of residual stocking. In the longer term, residual stocking had a strong influence on leaf area increase per tree and was correlated with changes in crown length.     

Sources of within- and between-stand variability in specific leaf area of three ecologically distinct conifer species

Specific leaf area (SLA) is an important ecophysiological variable, but its variability within and between stands has rarely been simultaneously examined and modeled across multiple species. Extensive datasets on SLA in coastal Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco), hybrid spruce (Picea engelmannii Parry × Picea glauca (Moench) Voss × Picea sitchensis (Bong.) Carr.), and ponderosa pine (Pinus ponderosa Dougl. ex P. & C. Laws.) were used to estimate variability of SLA within a canopy and its relationship to tree- and stand-level covariates, and to predict SLA at various locations in tree crowns. Also, in the case of hybrid spruce, variation in SLA due to different relative horizontal lengths from the bole was examined. In all species, SLA systematically increased from tree tip to crown base and decreased with foliage age class. Cardinal direction did not have a highly significant influence in either Douglas-fir or hybrid spruce, but SLA did significantly decrease from branch tip to bole in hybrid spruce. Tree- and stand-level (e.g. density, site index) factors had relatively little influence on SLA, but stand age did have a significant positive influence. For ponderosa pine, a significant relationship between canopy mean current-year SLA and carbon isotope discrimination was also found, suggesting the importance of water stress in this species. An equation was fitted to estimate SLA at various points in the canopy for each species and foliage age class using absolute height in the canopy, relative vertical height in the tree, and stand age.     

Leaf weight per area and leaf size of 85 Estonian woody species in relation to shade tolerance and light availability

Leaf weight per area (LWA) and leaf size were examined in 85 species of woody plants representing 83% of Estonian native flora.

Average values of 68.6 g m⁻² for LWA and 6.62 cm² for leaf size were estimated for the medium light availability. Mean LWA values can be used to correct for the bias in estimating foliage functional properties in forest stand models and for converting leaf biomass into foliage area. LWA increased with relative light availability (1 minus stand canopy cover) and species light demand, indicating that LWA was higher at the same light availability for more intolerant species. Leaf size is a canopy parameter that should be considered when canopy radiative balance and distribution of solar radiation is of interest. Mean leaf size was not so closely related to light availability as LWA, decreasing with species light demand and increasing with total plant height. Variation in leaf size is discussed in terms of species shade tolerance and height growth.     

Influence of light availability on growth, leaf morphology and plant architecture of beech ( Fagus sylvatica L.), maple ( Acer pseudoplatanus L.) and ash ( Fraxinus excelsior L.) saplings

In a field study, we measured saplings of beech, ash and maple growing in a fairly even-aged mixed-species thicket established by natural regeneration beneath a patchy shelterwood canopy with 3–60% of above canopy radiation reaching the saplings. Under low light conditions, maple and ash showed a slight lead in recent annual length increment compared with beech. With increasing light, ash and maple constantly gained superiority in length increment, whereas beech approached an asymptotic value above 35% light. A suite of architectural and leaf morphological attributes indicated a more pronounced ability of beech to adapt to shade than ash and maple. Beech displayed its leaves along the entire tree height (with a concentration in the middle crown), yielding a higher live crown ratio than ash and maple. It allocated biomass preferentially to radial growth which resulted in low height to diameter ratios, and expressed marked plagiotropic growth in shade indicating a horizontal light-foraging strategy. In addition, beech exhibited the highest specific leaf area, a greater total leaf area per unit tree height, a slightly greater leaf area index, and a greater plasticity to light in total leaf area. Ash and maple presented a “gap species” growth strategy, characterized by a marked and constant response in growth rates to increasing light and an inability to strongly reduce their growth rates in deep shade. In shade, they showed some plasticity in displaying most of their leaf area at the top of the crown to minimize self-shading and to enhance light interception. Through this, particularly, maple developed an “umbrella” like crown. These species-specific responses may be used for controlling the development of mixed-species regeneration in shelterwood systems.     

Effects of crown development on leaf irradiance,leaf morphology and photosynthetic capacity in a peach tree

The three-dimensional (3-D) architecture of a peach tree (Prunus persica L. Batsch) growing in an orchard near Avignon, France, was digitized in April 1999 and again four weeks later in May 1999 to quantify increases in leaf area and crown volume as shoots developed. A 3-D model of radiation transfer was used to determine effects of changes in leaf area density and canopy volume on the spatial distribution of absorbed quantum irradiance (PAR(a)). Effects of changes in PAR(a) on leaf morphological and physiological properties were determined. Leaf mass per unit area (M(a)) and leaf nitrogen concentration per unit leaf area (N(a)) were both nonlinearly related to PAR(a), and there was a weak linear relationship between leaf nitrogen concentration per unit leaf mass (N(m)) and PAR(a). Photosynthetic capacity, defined as maximal rates of ribulose-1,5-bisphosphate carboxylase (Rubisco) carboxylation (V(cmax)) and electron transport (J(max)), was measured on leaf samples representing sunlit and shaded micro-environments at the same time that the tree crown was digitized. Both V(cmax) and J(max) were linearly related to N(a) during May, but not in April when the range of N(a) was low. Photosynthetic capacity per unit N(a) appeared to decline between April and May. Variability in leaf nitrogen partitioning between Rubisco carboxylation and electron transport was small, and the partitioning coefficients were unrelated to N(a). Spatial variability in photosynthetic capacity resulted from acclimation to varying PAR(a) as the crown developed, and acclimation was driven principally by changes in M(a) rather than the amount or partitioning of leaf nitrogen.     

Shoot biomass growth is related to the vertical leaf nitrogen gradient in Salix canopies

Plant canopy optimization models predict that leaf nitrogen (N) distribution in the canopy will parallel the vertical light gradient, and numerous studies with many species have confirmed this prediction. Further, it is predicted that for a given canopy leaf area, a low vertical light extinction coefficient will promote rapid growth. Therefore, the ideal canopy of fast-growing plants should combine high leaf area index with a low light extinction coefficient; the latter being reflected in a flat vertical leaf N gradient throughout the canopy. Based on data from an experimental Salix stand (six varieties) grown on agricultural land in central Sweden, we tested the hypothesis that shoot growth is correlated with vertical leaf N gradient in canopies of hybrid willows bred for biomass production, which could have implications for Salix breeding. Tree improvement research requires screening of growth-related traits in large numbers of plants, but assessment of canopy leaf N gradients by chemical analysis is expensive, time-consuming and destructive. An alternative to analytical methods is to estimate leaf N gradients nondestructively with an optical chlorophyll meter (SPAD method). Here we provide a specific calibration for interpreting SPAD data measured in hybrid willows grown in biomass plantations on fertile agricultural land. Based on SPAD measurements, a significant and inverse relationship (r(2) = 0.88) was found between shoot biomass growth and vertical leaf N gradient across canopies of six Salix varieties.     

Canopy development in young sugi (Cryptomeria japonica) stands in relation to changes with age in crown morphology and structure

In young sugi (Cryptomeria japonica D. Don) stands, crown shape (crown length/crown diameter) ratio, average branch inclination, and spatial density of foliage in the crown increased with stand age. Within crowns, foliage distribution increased from the apex downward and, until crown closure, reached a maximum near the crown base. After crown closure, the maximum occurred near the middle of the crown. In each stand, foliage distribution in the canopy showed almost the same vertical change over time as it did in individual crowns. The vertical distribution of foliage in the canopy moved upward with stand age, accompanied by an increase in canopy depth and leaf mass. The shape of the vertical distribution was almost symmetrical between the upper and lower halves in the closed stands, although slightly skewed downward. The logarithm of average spatial density decreased linearly as cumulative leaf mass increased with distance from the top of the canopy. The total cross-sectional area of the crowns exceeded the stand area from the middle of the canopy downward in the closed stands because of crown overlap. However, partly because of changes in crown morphology and structure, the increase in leaf mass with stand age did not always cause more severe crown competition.     

Coordination of crown structure, leaf plasticity and carbon gain within the crowns of three winter-deciduous mature trees

We examined the vertical profiles of leaf characteristics within the crowns of two late-successional (Fagus crenata Blume and Fagus japonica Maxim.) and one early-successional tree species (Betula grossa Sieb. et Zucc.) in a Japanese forest. We also assessed the contributions of the leaves in each crown layer to whole-crown instantaneous carbon gain at midday. Carbon gain was estimated from the relationship between electron transport and photosynthetic rates. We hypothesized that more irradiance can penetrate into the middle of the crown if the upper crown layers have steep leaf inclination angles. We found that such a crown has a high whole-crown carbon gain, even if leaf traits do not change greatly with decreasing crown height. Leaf area indices (LAIs) of the two Fagus trees (5.26-5.52) were higher than the LAI of the B. grossa tree (4.50) and the leaves of the F. crenata tree were more concentrated in the top crown layers than were leaves of the other trees. Whole-crown carbon gain per unit ground area (micromol m(-2) ground s(-1)) at midday on fine days in summer was 16.3 for F. crenata, 11.0 for F. japonica, and 20.4 for B. grossa. In all study trees, leaf dry mass (LMA) and leaf nitrogen content (N) per unit area decreased with decreasing height in the crown, but leaf N per unit mass increased. Variations (plasticity) between the uppermost and lowermost crown layers in LMA, leaf N, the ratio of chlorophyll to N and the ratio of chlorophyll a to b were smaller for F. japonica and B. grossa than for F. crenata. The light extinction coefficients in the crowns were lower for the F. japonica and B. grossa trees than for the F. crenata tree. The leaf carbon isotope ratio (delta(13)C) was higher for F. japonica and B. grossa than for F. crenata, especially in the mid-crown. These results suggest that, in crowns with low leaf plasticity but steep leaf inclination angles, such as those of F. japonica and B. grossa trees, irradiance can penetrate into the middle of the crowns, thereby enhancing whole-crown carbon gain.     

Structural differences and functional similarities between two sugar maple (Acer saccharum) stands

The spatially inexplicit or functional multilayer models used to predict canopy transpiration or photosynthesis are based on the assumption that closed stands show less functional variability than structural variability, because foliage tends to arrange itself in space to optimize the capture of light. To validate this assumption, we compared the structural and functional properties, and the measured and modeled transpiration fluxes of two sugar maple (Acer saccharum Marsh.) stands of comparable leaf mass but differing in height and diameter distributions. One stand was characterized by a well-developed single-layer canopy, whereas the other stand had a multilayered canopy and a stem diameter distribution of the classical inverse-J shape. Stand differences in height and diameter distribution, and canopy gap fraction, were highly significant. There were minor but significant differences in leaf mass and leaf mass per unit leaf area (LMA) distributions. We found no differences in tree-level relationships between basal area and either transpiration flux or sapwood area. We compared measurements of stand transpiration with transpiration estimates obtained from a multilayer gas exchange model, in which only the nonspatial inputs, leaf area index and LMA frequency distribution described stand structure. For both stands, modeled values of daily transpiration closely followed measured values (r(2) = 0.94). These results support use of the nonspatially explicit approach to estimating canopy gas exchange, especially if the intent is to scale-up to larger portions of the landscape.     

Variability in Populus leaf anatomy and morphology in relation to canopy position, biomass production, and varietal taxon

Twelve poplar (Populus) genotypes, belonging to different taxa and to the sections Aigeiros and Tacamahaca, were studied during the third growing season of the second rotation of a high density coppice culture. With the objective to highlight the relationships between leaf traits, biomass production and taxon as well as the influence of canopy position, anatomical and morphological leaf characteristics (i.e. thickness of epidermis, of palisade and spongy parenchyma layers, density and length of stomata, leaf area, specific leaf area (SLA) and nitrogen concentration) were examined for mature leaves from all genotypes and at two canopy positions (upper and lower canopy). Above ground biomass production, anatomical traits, stomatal and morphological leaf characteristics varied significantly among genotypes and between canopy positions. The spongy parenchyma layer was thicker than the palisade parenchyma layer for all genotypes and irrespective of canopy position, except for genotypes belonging to the P. deltoides × P. nigra taxon (section Aigeiros). Leaves at the upper canopy position had higher stomatal density and thicker anatomical layers than leaves at the lower canopy position. Leaf area and nitrogen concentration increased from the bottom to the top of the canopy, while SLA decreased. Positive correlations between biomass production and abaxial stomatal density, as well as between biomass production and nitrogen concentration were found. A principal component analysis (PCA) showed that genotypes belonging to the same taxon had similar anatomical characteristics, and genotypes of the same section also showed common leaf characteristics. However, Wolterson (P. nigra) differed in anatomical leaf characteristics from other genotypes belonging to the same section (section Aigeiros). Hybrids between the two sections (Aigeiros × Tacamahaca) expressed leaf characteristics intermediate between both sections, while their biomass production was low.     

Petiole length and biomass investment in support modify light interception efficiency in dense poplar plantations

Leaf architecture, stand leaf area index and canopy light interception were studied in 13 poplar clones growing in a second rotation of a coppice plantation, to determine the role of leaf architectural attributes on canopy light-harvesting efficiency and to assess biomass investment in leaf support tissue. Stand leaf area index (L) varied from 2.89 to 6.99, but L was only weakly associated with canopy transmittance (TC). The weak relationship between TC and L was a result of a higher degree of foliage aggregation at larger values of L, leading to lower light-interception efficiency in stands with greater total leaf area. We observed a strong increase in leaf aggregation and a decrease in light-harvesting efficiency with decreasing mean leaf petiole length (PL) but not with leaf size, possibly because, in cordate or deltoid poplar leaves, most of the leaf area is located close to the petiole attachment to the lamina. Although PL was the key leaf characteristic of light-harvesting efficiency, clones with longer petioles had larger biomass investments in petioles, and there was a negative relationship between PL and L, demonstrating that enhanced light harvesting may lead to an overall decline in photosynthesizing leaf surface. Upper-canopy leaves were generally larger and had greater dry mass (MA) and nitrogen per unit area (NA) than lower-canopy leaves. Canopy plasticity in MA and NA was higher in clones with higher foliar biomass investment in midrib, and lower in clones with relatively longer petioles. These relationships suggest that there is a trade-off between photosynthetic plasticity and biomass investment in support, and also that high light-harvesting efficiency may be associated with lower photosynthetic plasticity. Our results demonstrate important clonal differences in leaf aggregation that are linked to leaf structure and biomass allocation patterns within the leaf.