INTERACTIONS BETWEEN CROWN STRUCTURE AND LIGHT ENVIRONMENT IN FIVE RAIN FOREST PIPER SPECIES

Measurements of light variation among leaves within crowns of five Piper species were compared with estimates of spatial variation in light within understory, forest edge, and clearing habitats to estimate the extent to which crown structure contributes to variation in leaf light environment. Daily photon flux density (PFD) varied greatly within and among crowns. Coefficients of variation for daily PFD among sensors within a single crown ranged from 26 to 79%. Within a single crown located in a clearing, the range in daily PFD among leaves was nearly as great as the range over the entire sample of plants. In the understory, localized sunfleck activity contributed to a high degree of spatial variation in instantaneous and total PFD among leaves within individual crowns. Much of the microsite variation in sunfleck activity, however, reflected environmental conditions within the understory habitat. Within an array of sensors placed next to Piper crowns in the understory, correlations were poor for light sensors spaced only 0.2 m apart, and only 8% of the variance in light readings was explained by measurements made 0.5 m away. In the clearing habitats, microsite heterogeneity among leaves was more strongly influenced by leaf positions within crowns and leaf angles than by spatial heterogeneity within the habitat.     

Comparison of the physiology, morphology, and leaf demography of tropical saplings with different crown shapes

Branch architecture, leaf photosynthetic traits, and leaf demography were investigated in saplings of two woody species, Homolanthus caloneurus and Macaranga rostulata, co-occurring in the understory of a tropical mountain forest. M. rostulata saplings have cylindrical crowns, whereas H. caloneurus saplings have flat crowns. Saplings of the two species were found not to differ in area-based photosynthetic traits and in average light conditions in the understory of the studied site, but they do differ in internode length, leaf emergence rate, leaf lifespan, and total leaf area. Displayed leaf area of H. caloneurus saplings, which have the more rapid leaf emergence, was smaller than that of M. rostulata saplings, which have a longer leaf lifespan and larger total leaf area, although M. rostulata saplings showed a higher degree of leaf overlap. Short leaf lifespan and consequent small total leaf area would be linked to leaf overlap avoidance in the densely packed flat H. caloneurus crown. In contrast, M. rostulata saplings maintained a large total leaf area by producing leaves with a long leaf lifespan. In these understory saplings with a different crown architecture, we observed two contrasting adaptation strategies to shade which are achieved by adjusting a suite of morphological and leaf demographic characters. Each understory species has a suite of morphological traits and leaf demography specific to its architecture, thus attaining leaf overlap avoidance or large total leaf area.     

Branch architecture,light interception and crown development in saplings of a plagiotropically branching tropical tree,Polyalthia jenkinsii (Annonaceae)

To investigate crown development patterns, branch architecture, branch-level light interception, and leaf and branch dynamics were studied in saplings of a plagiotropically branching tree species, Polyalthia jenkinsii Hk. f. & Thoms. (Annonaceae) in a Malaysian rain forest. Lengths of branches and parts of the branches lacking leaves ('bare' branches) were smaller in upper branches than in lower branches within crowns, whereas lengths of 'leafy' parts and the number of leaves per branch were larger in intermediate than in upper and lower branches. Maximum diffuse light absorption (DLA) of individual leaves was not related to sapling height or branch position within crowns, whereas minimum DLA was lower in tall saplings. Accordingly, branch-level light interception was higher in intermediate than in upper and lower branches. The leaf production rate was higher and leaf loss rate was smaller in upper than in intermediate and lower branches. Moreover, the branch production rate of new first-order branches was larger in the upper crowns. Thus, leaf and branch dynamics do not correspond to branch-level light interception in the different canopy zones. As a result of architectural constraints, branches at different vertical positions experience predictable light microenvironments in plagiotropic species. Accordingly, this pattern of carbon allocation among branches might be particularly important for growth and crown development in plagiotropic species.     

A tropical epiphytic orchid uses a low‐light interception strategy in a spatially heterogeneous light environment

Light is considered a non‐limiting factor for vascular epiphytes. Nevertheless, an epiphyte's access to light may be limited by phorophyte shading and the spatio‐temporal environmental patchiness characteristic of epiphytic habitats. We assessed the extent to which potential light interception in Rodriguezia granadensis, an epiphytic orchid, is determined by individual factors (plant size traits and leaf traits), or environmental heterogeneity (light patchiness) within the crown of the phorophyte, or both. We studied 104 adult plants growing on Psidium guajava trees in two habitats with contrasting canopy cover: a dry tropical forest edge, and isolated trees in a pasture. We recorded the number of leaves and the leaf area, the leaf position angles, and the potential exposure of the leaf surface to direct irradiance (silhouette area of the leaf blade), and the potential irradiance incident on each plant. We found the epiphytes experience a highly heterogeneous light environment in the crowns of P. guajava. Nonetheless, R. granadensis plants displayed a common light interception strategy typical of low‐light environments, resembling terrestrial, forest understory plants. Potential exposure of the total leaf surface to direct irradiance correlated positively with plant size and within‐plant variation in leaf orientation. In many‐leaved individuals, within‐plant variation in leaf angles produced complementary leaf positions that enhanced potential light interception. This light interception strategy suggests that, in contrast to current wisdom, enhancing light capture is important for vascular epiphytes in canopies with high spatio‐temporal heterogeneity in light environments.     

Implications of opposite phyllotaxis for light interception efficiency of Mediterranean woody plants

Opposite leaves lead to a greater leaf overlapping than leaves spirally arranged along a shoot, decreasing light interception efficiency (Ea, fraction of the light reaching the plant actually intercepted by the leaves) of the crown. However, Ea results from a whole suite of morphological traits. The interplay between phyllotaxis, crown architecture, leaf morphology and Ea was explored in 12 woody species from Mediterranean-type ecosystems, where the abundance of woody species with opposite phyllotaxis is unusually high. The three-dimensional model Y-plant was used to estimate Ea in unbranched, vertical shoots of each species encompassing the natural morphological variation found from moderate shade to open light environments. Ea exhibited significant interspecific differences, ranging from 0.25 in Daphne gnidium to 0.75 in Cistus ladanifer, Olea europaea and Salvia officinalis, decreasing with leaf inclination angle and leaf area ratio (LAR), and increasing with internode-to-leaf-length ratio and supporting biomass. Species with spiral vs. opposite phyllotaxis did not differ in their mean Ea. However, the former had higher Ea than the latter at short internode lengths. The natural range of variation in internode length had a larger effect on Ea than the natural range of leaf elevation angle. Principal component analysis segregated species with opposite phyllotaxis from those with spiral leaves because of their greater self-shading for high sun elevation angles (>45°); they were in turn distributed in two groups, one with high Ea, large investment in supporting biomass and long internodes, and another with low Ea and large LAR. Species with spiral phyllotaxis all had intermediate or low Ea and steep leaf elevation angles. Species with opposite phyllotaxis can compensate their less efficient leaf arrangement by decreasing leaf elevation angle and increasing internode length, but they may experience a real phylogenetic constraint for light interception when biomass allocation to supporting tissues (internodes and petioles) becomes very costly. This constraint could be involved in the shade intolerance of woody Mediterranean species exhibiting opposite phyllotaxis.     

Light interception efficiency explained by two simple variables: a test using a diversity of small- to medium-sized woody plants

? Plant light interception efficiency is a crucial determinant of carbon uptake by individual plants and by vegetation. Our aim was to identify whole-plant variables that summarize complex crown architecture, which can be used to predict light interception efficiency. ? We gathered the largest database of digitized plants to date (1831 plants of 124 species), and estimated a measure of light interception efficiency with a detailed three-dimensional model. Light interception efficiency was defined as the ratio of the hemispherically averaged displayed to total leaf area. A simple model was developed that uses only two variables, crown density (the ratio of leaf area to total crown surface area) and leaf dispersion (a measure of the degree of aggregation of leaves). ? The model explained 85% of variation in the observed light interception efficiency across the digitized plants. Both whole-plant variables varied across species, with differences in leaf dispersion related to leaf size. Within species, light interception efficiency decreased with total leaf number. This was a result of changes in leaf dispersion, while crown density remained constant. ? These results provide the basis for a more general understanding of the role of plant architecture in determining the efficiency of light harvesting.     

How is light interception efficiency related to shoot structure in tall canopy species?

Coexistence of multiple species is a fundamental aspect of plant and forest ecology. Although spatial arrangement of leaves within crowns is an important determinant of light interception and productivity, shoot structure varies considerably among coexisting canopy species. We investigated the relative importance of structural traits in determining the light availability of leaves (I) and light interception efficiency at the current-year shoot level (LIE_CS; the total light interception of leaves divided by shoot biomass) at the top of crowns of 11 canopy species in a cool-temperate forest in Japan. In accordance with Corner’s rules, the total mass, stem mass, total mass of leaf laminae, individual leaf area, and stem cross-sectional area of current-year shoot were positively correlated with each other, and branching intensity (the number of current-year shoots per branch unit of 1-m length) was inversely correlated with these traits across species. In contrast, I was correlated not with these traits, but with leaf elevation angle (a _L). Moreover, variation in LIE_CS across species was caused by variation in I (thus in a _L). Thus, a _L is a key parameter for the leaf light interception of canopy shoots in this cool-temperate forest. Differences in a _L across species might be related to different physiological strategies that developed in the high light and water-limited environment of forest canopies. Small variation in the length of current-year shoots among species implies that variations in I and LIE_CS would be important for the coexistence of these canopy species.     

Above-ground biomass investments and light interception of tropical forest trees and lianas early in succession

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. METHODS: 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 Phi(d) was calculated with a canopy model. The model was then used to estimate light interception per unit of leaf mass (Phi(leaf mass)), total above-ground mass (Phi(mass)) and crown structure efficiency (E(a), the ratio of absorbed vs. available light). KEY RESULTS: The SLPs Trema and Ochroma intercepted higher amounts of light per unit leaf mass (Phi(leaf mass)) because they had shallower crowns, resulting in higher crown use efficiency (E(a)) than the other species. These SLPs (but not Cecropia) were also taller and intercepted more light per unit leaf area (Phi(area)). LLPs and lianas had considerably higher amounts of leaf mass and area per unit above-ground mass (LMR and LAR, respectively) and thus attained Phi(mass) values similar to the SLPs (Phi(mass)=Phi(area)xLAR). Lianas, which were mostly self-supporting, had light interception efficiencies similar to those of the trees. CONCLUSIONS: 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.     

Irradiance heterogeneity within crown affects photosynthetic capacity and nitrogen distribution of leaves in Cedrela sinensis

Because light conditions in the forest understory are highly heterogeneous, photosynthetic acclimation to spatially variable irradiance within a crown is important for crown‐level carbon assimilation. The effect of variation in irradiance within the crown on leaf nitrogen content and photosynthetic rate was examined for pinnate compound leaves in saplings of Cedrela sinensis, a pioneer deciduous tree. Five shading treatments, in which 0, 25, 50, 75 and 100% of leaves were shaded, were established by artificial heavy shading using shade screen umbrellas with 25% transmittance. Although the nitrogen content of leaves was constant regardless of shading treatment, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) content and light‐saturated photosynthetic capacity were lower in shade leaves within partially shaded crowns than within fully shaded crowns. Shade leaves within partially shaded crowns contained higher amount of amino acids. Most shade leaves died in partially shaded crowns, whereas more than half of shade leaves survived in totally shaded crowns. Assumptions on photosynthetic acclimation to local light conditions cannot explain why shade leaves have different photosynthetic capacities and survival rates in between partially and totally shaded crowns. Irradiance heterogeneity within the crown causes a distinct variation in photosynthetic activity between sun and shaded leaves within the crown.     

PLANT SIZE AND FORM IN THE UNDERSTORY PALM GENUS GEONOMA: ARE SPECIES VARIATIONS ON A THEME?

This study addressed the hypothesis that phylogenetic changes in plant size at reproductive maturity may have facilitated adaptive radiation of Geonoma species within rain forest understory habitats. Leaf size, leaf form, plant size, and growth form were compared within and among 23 species of Geonoma from lowland and montane rain forest areas of Costa Rica and Colombia. Leaf size was significantly correlated with crown height in 18 of the 21 species examined, and with stem diameter in 17 of the species. In species characterized by a gradual ontogenetic transition from bifid to dissected leaves, shoots with bifid leaves were significantly smaller than shoots with dissected leaves with respect to rachis length, number of plications, and stem diameter. Among species, stem diameter below the crown explained 74% of the variation in leaf size (rachis length). Crown height and stem diameter were positively correlated among clustered species, but not among solitary species or all species combined. Leaf dissection was correlated with crown height among the 17 species with dissected leaves; species with bifid leaves were significantly smaller than species with dissected leaves with respect to leaf size and stem diameter. Solitary species had larger leaves and larger stem diameters than clustered species at the same crown heights. Morphological patterns among species generally followed within-species trends. These patterns suggest that Geonoma species are variants on a generic theme:within and among species, leaf size and complexity of form increase with stem diameter and crown height. Solitary and clustered growth forms appear to be morphologically convergent; within each of these architectural groups, the generic theme still applies. Evolutionary changes in leaf size, leaf form, and plant size, however, have clearly involved other factors in addition to variation in plant size.     

The role of crown architecture, leaf phenology and photosynthetic activity in promoting complementary use of light among coexisting species in temperate forests

Mixed forests comprising multiple tree species with contrasting crown architectures, leaf phenologies, and photosynthetic activity, tend to have high ecosystem productivity. We propose that in such forests, differentiation among coexisting species in their spatial and temporal strategies for light interception, results in complementary use of light. Spatial differentiation among coexisting tree species occurs as a result of adaptation of crown architecture and shoot/leaf morphology to the spatially variable light conditions of the canopy, sub-canopy, and understory. Temporal differentiation occurs as a result of variation in leaf phenology and photosynthetic activity. The arrangement of leaves in both space and time is an important aspect of plant strategies for light interception and determines photosynthetic carbon gain of the plant canopy. For example, at the shoot level, morphological and phenological differentiation between long and short shoots reflects their respective shoot functions, indicating that spatial and temporal strategies for light interception are linked. Complementary use of light is a consequence of the spatiotemporal differentiation in light interception among coexisting species. Because coexisting species may show differentiation in strategies for resource acquisition (functional diversification) or convergence with respect to some limiting resource (functional convergence), the relative importance of various crown functions and their contribution to growth and survival of individuals need to be evaluated quantitatively and compared among coexisting species.     

Leaf position,light levels,and nitrogen allocation in five species of rain forest pioneer trees

We used path analysis to ask whether leaf position or leaf light level was a better predictor of within-plant variation in leaf nitrogen concentration in five species of rain forest pioneer trees (Cecropia obtusifolia, Ficus insipida, Heliocarpus appendiculatus, Piper auritum, and Urera caracasana) from the Los Tuxtlas Biological Station, Veracruz, Mexico. Three hundred seventy-five leaves on 28 plants of the five species were analyzed for leaf nitrogen concentration, leaf mass per area, and leaf light interception at different positions (= nodes) along a shoot. Mean values of leaf nitrogen concentration ranged from 0.697 to 0.993 g/m² in the five species, and varied by as much as 2.24 g/m² among leaves on individual plants. Leaf position on the shoot explained significantly more of the within-plant variation in leaf nitrogen concentration than did leaf light level in four of the five species: Cecropia obtusifolia, Heliocarpus appendiculatus, Piper auritum (branch leaves only), and Urera caracasana. However, individual species differed considerably in the patterns of nitrogen allocation and leaf mass per area among leaves on a shoot. These results suggest that leaf nitrogen deployment in these plants is, in part, developmentally constrained and related to the predictability of canopy light distribution associated with plant growth form.     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

遮荫处理对梅叶冬青叶片形态、光合特性和生长的影响

为了解遮荫环境对梅叶冬青(Ilex asprella)生长和光合特性的影响,采用遮荫网的方法模拟85%、56%和全光照等3种光照环境,研究了遮荫对其生长、光合参数以及生物量等的影响。结果表明,经一年遮荫处理后,梅叶冬青的叶绿素a、b和叶绿素总量都随着遮荫强度的增加而显著增高,胡萝卜素含量则显著降低。与对照相比,56%遮荫处理显著提高梅叶冬青的最大净光合速率和光饱和点,分别提高了17.6%和25.2%,但是85%遮阴处理则显著降低最大净光合速率和光饱和点,分别降低了18.2%和24.1%,两种遮荫处理均显著降低了光补偿点。叶长、叶宽、比叶面积、单片叶面积和叶片含水量均随着遮荫强度的增加而显著增加,而叶片厚度则显著减小。遮荫处理明显抑制整株生物量增长,减小根冠比,但是株高、冠幅和径向生长随遮荫处理时间不同而有所变异。因此,梅叶冬青有耐荫偏阳的特性,在林下种植时需及时调控乔冠层的透光率,一般应大于44%。     

A forest-structure-based analysis of rain flow into soil in a dense deciduous Betula ermanii forest with understory dwarf bamboo

The relationship between rain flow into the soil and forest structure was investigated in a dense deciduous Betula ermanii forest in northern Japan. The forest floor was covered with dwarf bamboo Sasa kurilensis. Observation was conducted from mid-July to late October in 1998. Leaf fall of Betula started in early September and ended in late October. Stemflow was proportional to rainfall and tree size [diameter at breast height (DBH)], and for the same rainfall, stemflow increased with leaf fall. On the contrary, throughfall decreased with leaf fall. Throughfall was intercepted also by Sasa in proportion to its leaf area. Multiple linear regression analysis revealed that stemflow and throughfall of Betula and Sasa were predictable as functions of rainfall and forest structural characteristics, such as DBH, tree density, and stand leaf mass. The rain interception by plants tended to decrease from summer to autumn, but the difference in the interception was about 2% between July (fully expanded leaves) and late October (lack of leaves). About 96 and 87% of rainfall reached the above- and below-Sasa layers, respectively. Thus, this study showed that understory Sasa is a major component of rain interception within the stand and that rain flow into the soil can be estimated by using rainfall and the forest structural parameters, such as DBH, tree density and stand leaf mass.     

The expression of light-related leaf functional traits depends on the location of individual leaves within the crown of isolated Olea europaea trees

Background The spatial arrangement and expression of foliar syndromes within tree crowns can reflect the coupling between crown form and function in a given environment. Isolated trees subjected to high irradiance and concomitant stress may adjust leaf phenotypes to cope with environmental gradients that are heterogeneous in space and time within the tree crown. The distinct expression of leaf phenotypes among crown positions could lead to complementary patterns in light interception at the crown scale.Methods We quantified eight light-related leaf traits across 12 crown positions of ten isolated Olea europaea trees in the field. Specifically, we investigated whether the phenotypic expression of foliar traits differed among crown sectors and layers and five periods of the day from sunrise to sunset. We investigated the consequences in terms of the exposed area of the leaves at the tree scale during a single day.Key Results All traits differed among crown positions except the length-to-width ratio of the leaves. We found a strong complementarity in the patterns of the potential exposed area of the leaves among day periods as a result of a non-random distribution of leaf angles across the crown. Leaf exposure at the outer layer was below 60 % of the displayed surface, reaching maximum interception during morning periods. Daily interception increased towards the inner layer, achieving consecutive maximization from east to west positions within the crown, matching the sun’s trajectory.Conclusions The expression of leaf traits within isolated trees of O. europaea varies continuously through the crown in a gradient of leaf morphotypes and leaf angles depending on the exposure and location of individual leaves. The distribution of light-related traits within the crown and the complementarity in the potential exposure patterns of the leaves during the day challenges the assumption of low trait variability within individuals.     

Growth responses to arbuscular mycorrhizae by rain forest seedlings vary with light intensity and tree species

Light intensity and root colonization by arbuscular mycorrhizal (AM) fungi are considered important factors affecting the performance of rain forest plants, yet few studies have examined how these two factors interact. Whether AM colonization promoted growth or caused shifts in biomass allocation in seedlings of four species of Australian rain forest tree (Flindersia brayleana, Acmena resa, Cryptocarya mackinnoniana and Cryptocarya angulata), grown in a glasshouse under light conditions that mimicked the shaded understory (3% PAR) and small light gaps (10% PAR), was examined. Seedlings were grown in sterilized field soil and either inoculated with AM fungi or provided sterile inoculum. Four major findings emerged. First, in all species, seedlings grown in small gap light intensities were larger than seedlings grown in understory light intensities. Second, when seedling biomass was included as a covariate, variation in light intensity was associated with significant shifts in biomass allocation. In all species, leaf area ratio was lower at 10% PAR than at 3% PAR, while root-to-shoot ratio showed the opposite pattern in one of the four species (C. mackinonniana). Third, although percentage root length colonized by AM fungi was greater at 10% PAR than 3% PAR in all species, this difference could be accounted for by variation in seedling size in all species except C. angulata. Fourth, growth and biomass allocation responses to AM colonization varied with light intensity and plant species. AM colonization promoted growth in both light regimes only in F. brayleana, while it had no effect on growth in C. mackinnoniana and C. angulata in either light regime and promoted growth only under high light in A. resa. AM colonization had no effect on leaf area ratio or root-to-shoot ratio in any of the species, and significantly altered specific root length in only one of the four species (C. mackinnoniana). These findings suggest that rain forest seedlings are highly variable in their growth responses to AM colonization and that some of this variability is related to the light intensity of the environment. Given that seedlings may spend many years in the shaded understory, these differences among species could have important effects on long-term seedling performance and seedling community dynamics.     

Functional correlates of leaf demographic response to gap release in saplings of a shade-tolerant tree,<Emphasis Type="Italic"> Elateriospermum tapos</Emphasis>

For a shade-tolerant SE Asian tropical tree, Elateriospermum tapos (Euphorbiaceae), we studied field-established saplings in gaps and the shaded understory to test the hypothesis that differences in leaf demography and leaf life span under contrasting light regimes should be functionally correlated with architecture, self-shading and nitrogen distribution within the sapling crown. Rates of leaf production and net leaf gain were greater for saplings in gaps than those in the understory. Median leaf life span was approximately 26 months in the gap saplings, while it was estimated to be greater than 38 months in the understory saplings. Consequently, gap saplings had a greater standing leaf number and experienced greater degrees of self-shading than understory saplings. Light availability at individual leaves, estimated by a combination of canopy photos and a three-dimensional architecture model, were negatively correlated with leaf age in gap saplings but not so in understory saplings. Leaf nitrogen content per unit area (N_area) was influenced more by light availability than by leaf age in the gap saplings. In contrast, in understory saplings, N_area was neither correlated with light availability nor with leaf age, and did not decrease significantly before 38 months in leaf age. We conclude that saplings of this shade-tolerant species apparently prolong their leaf life span in the shaded understory through slower rates of leaf production, lower standing number of leaves and lower degrees of self shading than in gap, and that the rate of decline of N_area with leaf age depends on architecture and self-shading regimes that respond to changes in light regimes.