Seedlings of temperate rainforest conifer and angiosperm trees differ in leaf area display

Background and Aims

The contemporary relegation of conifers mainly to cold or infertile sites has been ascribed to low competitive ability, as a result of the hydraulic inefficiency of tracheids and their seedlings'' initial dependence on small foliage areas. Here it is hypothesized that, in temperate rainforests, the larger leaves of angiosperms also reduce self-shading and thus enable display of larger effective foliage areas than the numerous small leaves of conifers.

Methods

This hypothesis was tested using 3-D modelling of plant architecture and structural equation modelling to compare self-shading and light interception potential of seedlings of six conifers and 12 angiosperm trees from temperate rainforests. The ratio of displayed leaf area to plant mass (LAR_d) was used to indicate plant light interception potential: LAR_d is the product of specific leaf area, leaf mass fraction, self-shading and leaf angle.

Results

Angiosperm seedlings self-shaded less than conifers, mainly because of differences in leaf number (more than leaf size), and on average their LAR_d was about twice that of conifers. Although specific leaf area was the most pervasive influence on LAR_d, differences in self-shading also significantly influenced LAR_d of large seedlings.

Conclusions

The ability to deploy foliage in relatively few, large leaves is advantageous in minimizing self-shading and enhancing seedling light interception potential per unit of plant biomass. This study adds significantly to evidence that vegetative traits may be at least as important as reproductive innovations in explaining the success of angiosperms in productive environments where vegetation is structured by light competition.     

Shoot morphology of Aucuba japonica incurred by anisophylly: ecological implications

Anisophylly, having leaves different in size and/or shape, was quantified in adult Aucuba japonica and simulations were carried out to evaluate the effects of anisophylly on the extent of self-shading at the single-shoot level as well as at the whole-canopy level. Clear anisophylly was observed in the individual after switching from the single-stemmed juvenile stage to the multi-stemmed adult stage. In such plants, leaf area in the canopy abruptly increased. The effective display of adult foliage involved a variety of morphological changes in addition to anisophylly, most prominently reduction in leaf size compared to juveniles. The simulation results indicate that diversity of leaf size and shape is an effective means of minimizing self-shading as well as allowing the efficient exploitation of a larger canopy volume in adult plants. Anisophylly also increased the biomass use efficiency of individual plants at maturity. Taken together, having diverse leaf forms is superior to having a single leaf form for maximizing area acquisition and for efficiently filling the acquired area. We therefore conclude that the anisophylly expressed in A. japonica is adaptive.     

The functional morphology of light capture and carbon gain in the Redwood forest understorey plant Adenocaulon bicolor Hook

1. A three-dimensional geometric simulation model of crown architecture was utilized to investigate the efficiency of light capture and its relationship to whole-plant CO₂ assimilation of Adenocaulon bicolor .
2. Positioning of the leaves by the combined effects of ontogenetic variations in petiole length and angle and leaf size, and the leaf divergence angles were shown to be effective in minimizing self shading. The efficiency of light absorption varied from 0·64 to 0·70 among individual plants that were sampled.
3. Plant to plant variation in simulated daily carbon gain was strongly influenced by variations in the direct and diffuse PFD received by the individual plants. When simulations were run for all plants under a single common light environment, the carbon gain was strongly dependent on the efficiencies of light absorption of the different plants.
4. Simulations in which petiole length was varied showed a non-linear dependence of light absorption efficiency on petiole length. When both petiole length and leaf size were varied in a way that maintained a constant biomass then an optimal petiole length that corresponded to the observed petiole length was apparent. The observed divergence angle between successive leaves also maximized light absorption efficiency as compared to greater or lesser angles, but increases in internode length had no significant effect.
5. The results of this study provide evidence for selection for an 'optimal design' of crown architecture in Adenocaulon bicolor that maximizes light capture.     

3D Architectural Modelling of Aerial Photomorphogenesis in White Clover (Trifolium repens L.) using L-systems

The objective of this work was to construct a model of aerialdevelopment of clover that takes into account morphogeneticresponses to the light environment, and to use it to analyseand understand these processes in terms of signal perceptionand integration. The plant model was interfaced with a MonteCarlo model that determines photosynthetically active radiation(PAR) and red/far-red ratio (R/FR) throughout the canopy, takinginto account the absorption, reflection and transmission oflight by individual leaves. Light intensity and quality weresensed by the plant model at discrete time intervals and atdiscrete sites of perception: apices, emerging internodes andpetiole tips. This input regulated the final size of internodesand leaves, the vertical positioning of leaves, and the branchingdelay. The empirical relations (regression functions) quantifyingthis regulation were derived from data reported in the literatureand original measurements. Simulations produced realistic visualizationsand quantitative characterizations of the modelled plants fordifferent light treatments. These results were in general agreementwith observations of real plants growing under similar conditions,suggesting that the dependence of organ size and position onlight treatments can be regarded as an integration of the responsesof individual plant organs to their local light environment.The model is used to describe the regulation of branch appearanceand the impact of self-shading on plant morphogenesis as a functionof local light environment. Copyright 2000 Annals of BotanyCompany Clover, Trifolium repens L, photomorphogenesis, plant architecture, L-system, modelling, Monte-Carlo method, competition for light, red : far-red ratio, irradiance, light quality, leaf size, self-shading     

Assessing transpiration in the tussock grass Stipa tenacissima L.: the crucial role of the interplay between morphology and physiology

Plant transpiration has a key role on both plant performance and ecosystem functioning in arid zones, but realistic estimates at appropriate spatial-temporal scales are scarce. Leaf and tiller morphology and crown architecture were studied together with leaf physiology and whole plant water balance in four individual plants of Stipa tenacissima of different sizes to determine the relative influence of processes taking place at different spatial and temporal scales on whole plant transpiration. Transpiration was estimated in potted plants by leaf-level gas exchange techniques (infrared gas analyzer and porometer), by sap flow measurements, and by integrating leaf physiology and crown architecture with the 3-D computer model Yplant. Daily transpiration of each individual plant was monitored using a gravimetric method, which rendered the reference values. Leaves on each individual plant significantly varied in their physiological status. Young and green parts of the leaves showed five times higher chlorophyll concentration and greater photosynthetic capacity than the senescent parts of the foliage. Instantaneous leaf-level transpiration measurements should not be used to estimate plant transpiration, owing to the fact that extrapolations overestimated individual transpiration by more than 100%. Considering leaf age effects and scaling the estimates according to the relative amount of each foliage category reduced this difference to 46% though it was still significantly higher than gravimetric measurements. Sap flow calculations also overestimated tussock transpiration. However, 3-D reconstruction of plants with Yplant and transpiration estimates, considering both the physiological status and the daily pattern of radiation experienced by each individual leaf section within the crown, matched the gravimetric measurements (differences were only 4.4%). The complex interplay of leaf physiology and crown structure must be taken into account in scaling up plant transpiration from instantaneous, leaf-level measurements, and our study indicates that transpiration of complex crowns is easily overestimated.     

Assessing transpiration in the tussock grass Stipa tenacissima L.: the crucial role of the interplay between morphology and physiology

Plant transpiration has a key role on both plant performance and ecosystem functioning in arid zones, but realistic estimates at appropriate spatial-temporal scales are scarce. Leaf and tiller morphology and crown architecture were studied together with leaf physiology and whole plant water balance in four individual plants of Stipa tenacissima of different sizes to determine the relative influence of processes taking place at different spatial and temporal scales on whole plant transpiration. Transpiration was estimated in potted plants by leaf-level gas exchange techniques (infrared gas analyzer and porometer), by sap flow measurements, and by integrating leaf physiology and crown architecture with the 3-D computer model Yplant. Daily transpiration of each individual plant was monitored using a gravimetric method, which rendered the reference values. Leaves on each individual plant significantly varied in their physiological status. Young and green parts of the leaves showed five times higher chlorophyll concentration and greater photosynthetic capacity than the senescent parts of the foliage. Instantaneous leaf-level transpiration measurements should not be used to estimate plant transpiration, owing to the fact that extrapolations overestimated individual transpiration by more than 100%. Considering leaf age effects and scaling the estimates according to the relative amount of each foliage category reduced this difference to 46% though it was still significantly higher than gravimetric measurements. Sap flow calculations also overestimated tussock transpiration. However, 3-D reconstruction of plants with Yplant and transpiration estimates, considering both the physiological status and the daily pattern of radiation experienced by each individual leaf section within the crown, matched the gravimetric measurements (differences were only 4.4%). The complex interplay of leaf physiology and crown structure must be taken into account in scaling up plant transpiration from instantaneous, leaf-level measurements, and our study indicates that transpiration of complex crowns is easily overestimated.     

Role of Leaf Surface Sugars in Colonization of Plants by Bacterial Epiphytes

The relationship between nutrients leached onto the leaf surface and the colonization of plants by bacteria was studied by measuring both the abundance of simple sugars and the growth of Pseudomonas fluorescens on individual bean leaves. Data obtained in this study indicate that the population size of epiphytic bacteria on plants under environmentally favorable conditions is limited by the abundance of carbon sources on the leaf surface. Sugars were depleted during the course of bacterial colonization of the leaf surface. However, about 20% of readily utilizable sugar, such as glucose, present initially remained on fully colonized leaves. The amounts of sugars on a population of apparently identical individual bean leaves before and after microbial colonization exhibited a similar right-hand-skewed distribution and varied by about 25-fold from leaf to leaf. Total bacterial population sizes on inoculated leaves under conditions favorable for bacterial growth also varied by about 29-fold and exhibited a right-hand-skewed distribution. The amounts of sugars on leaves of different plant species were directly correlated with the maximum bacterial population sizes that could be attained on those species. The capacity of bacteria to deplete leaf surface sugars varied greatly among plant species. Plants capable of supporting high bacterial population sizes were proportionally more depleted of leaf surface nutrients than plants with low epiphytic populations. Even in species with a high epiphytic bacterial population, a substantial amount of sugar remained after bacterial colonization. It is hypothesized that residual sugars on colonized leaves may not be physically accessible to the bacteria due to limitations in wettability and/or diffusion of nutrients across the leaf surface.     

Role of leaf surface sugars in colonization of plants by bacterial epiphytes

The relationship between nutrients leached onto the leaf surface and the colonization of plants by bacteria was studied by measuring both the abundance of simple sugars and the growth of Pseudomonas fluorescens on individual bean leaves. Data obtained in this study indicate that the population size of epiphytic bacteria on plants under environmentally favorable conditions is limited by the abundance of carbon sources on the leaf surface. Sugars were depleted during the course of bacterial colonization of the leaf surface. However, about 20% of readily utilizable sugar, such as glucose, present initially remained on fully colonized leaves. The amounts of sugars on a population of apparently identical individual bean leaves before and after microbial colonization exhibited a similar right-hand-skewed distribution and varied by about 25-fold from leaf to leaf. Total bacterial population sizes on inoculated leaves under conditions favorable for bacterial growth also varied by about 29-fold and exhibited a right-hand-skewed distribution. The amounts of sugars on leaves of different plant species were directly correlated with the maximum bacterial population sizes that could be attained on those species. The capacity of bacteria to deplete leaf surface sugars varied greatly among plant species. Plants capable of supporting high bacterial population sizes were proportionally more depleted of leaf surface nutrients than plants with low epiphytic populations. Even in species with a high epiphytic bacterial population, a substantial amount of sugar remained after bacterial colonization. It is hypothesized that residual sugars on colonized leaves may not be physically accessible to the bacteria due to limitations in wettability and/or diffusion of nutrients across the leaf surface.     

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.     

Ontogeny, understorey light interception and simulated carbon gain of juvenile rainforest evergreens differing in shade tolerance

Background and Aims

A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried.

Methods

The biomass distribution of juveniles (17–740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain.

Results

An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR_d = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey.

Conclusions

The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.     

Morphological variation of Aechmea distichantha (Bromeliaceae) in a Chaco forest: habitat and size-related effects

Plants show different morphologies when growing in different habitats, but they also vary in their morphology with plant size. We examined differences in sun- and shade-grown plants of the bromeliad Aechmea distichantha with respect to relationships between plant size and variables related to plant architecture, biomass allocation and tank water dynamics. We selected vegetative plants from the understorey and from forest edges of a Chaco forest, encompassing the whole size range of this bromeliad. Plant biomass was positively correlated with most architectural variables and negatively correlated with most biomass allocation variables. Understorey plants were taller and had larger diameters, whereas sun plants had more leaves, larger sheath area, sheath biomass and sheath mass fraction. All tank water-related variables were positively correlated with plant biomass. Understorey plants had a greater projected leaf area, whereas sun plants had higher water content and evaporative area. Plasticity indices were higher for water-related than for allocation variables. In conclusion, there were architectural and biomass allocation differences between sun- and shade-grown plants along a size gradient, which, in turn, affected tank water-related variables.     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

Responses of epidermal cell turgor pressure and photosynthetic activity of leaves of the atmospheric epiphyte Tillandsia usneoides (Bromeliaceae) after exposure to high humidity

It has been well-established that many epiphytic bromeliads of the atmospheric-type morphology, i.e., with leaf surfaces completely covered by large, overlapping, multicellular trichomes, are capable of absorbing water vapor from the atmosphere when air humidity increases. It is much less clear, however, whether this absorption of water vapor can hydrate the living cells of the leaves and, as a consequence, enhance physiological processes in such cells. The goal of this research was to determine if the absorption of atmospheric water vapor by the atmospheric epiphyte Tillandsia usneoides results in an increase in turgor pressure in leaf epidermal cells that subtend the large trichomes, and, by using chlorophyll fluorescence techniques, to determine if the absorption of atmospheric water vapor by leaves of this epiphyte results in increased photosynthetic activity. Results of measurements on living cells of attached leaves of this epiphytic bromeliad, using a pressure probe and of whole-shoot fluorescence imaging analyses clearly illustrated that the turgor pressure of leaf epidermal cells did not increase, and the photosynthetic activity of leaves did not increase, following exposure of the leaves to high humidity air. These results experimentally demonstrate, for the first time, that the absorption of water vapor following increases in atmospheric humidity in atmospheric epiphytic bromeliads is mostly likely a physical phenomenon resulting from hydration of non-living leaf structures, e.g., trichomes, and has no physiological significance for the plant's living tissues.     

Crown architecture in sun and shade environments: assessing function and trade-offs with a three-dimensional simulation model

Sun and shade environments place markedly different constraints on the photosynthetic performance of plants. Leaf-level photosynthetic responses to sun and shade have been extensively investigated, whereas there has been much less research on the functional role of crown architecture in these environments. This paper focuses on the role of architecture in maximizing light capture and photosynthesis in shaded understories and in minimizing exposure to excess radiation in open high light environments. Understanding these contrasting roles of architecture is facilitated by application of a three-dimensional structural-functional model, Y-plant. Surveys of understory plants reveal a diversity of architectures but a strong convergence at only modest light-capture efficiencies because of significant self-shading. Simulations with Psychotria species revealed that increasing internode lengths would increase light-capture efficiencies and whole plant carbon gain. However, the costs of the additional required biomechanical support was high, which, in terms of relative growth rates, would override the advantage provided by higher light-capture efficiencies. In high light environments, leaf angles and self-shading provide structural photoprotection, minimizing potential damage from photoinhbition. Simulations reveal that without these structural protections photoinhibition of photosynthesis is likely to be much greater with daily carbon gain significantly reduced.     

The study of a determinate growth orchid highlights the role of new leaf production in photosynthetic light acclimation

Plants usually respond to the changes of growth irradiance by a combination of the physiological modifications in their preexisting leaves and the production of new leaves. However, those with a determinate growth habit produce certain number of leaves in a growing season and cannot produce new leaves when light condition changes. We used an epiphytic orchid with only one leaf produced every growing season to examine whether and how determinate growth species adapt to changing environments after their preexisting leaves mature. Leaf photosynthesis and anatomy of Pleione aurita were investigated at full expansion and at 40 days after the fully expanded leaves were transferred from high to low light or from low to high light. Leaves show large physiological and morphological plasticity to light gradients at full expansion and the transferred leaves exhibited multiple physiological modifications, including reallocation of nitrogen between light harvesting and carbon fixation, and enhancement of thermal dissipation in their new environments, to optimize carbon assimilation and avoid photoinhibition. Irrespective of the various changes either to shade or sun, the sole preexisting leaf could not fully acclimate to new light environments due to the mesophyll thickness constraint. This leads to the consequence that only plants exposed to high light throughout the experiment had a positive annual biomass gain. Our results highlighted the importance of new leaf production in the carbon accumulation during photosynthetic light acclimation, and contribute new insights of epiphytes physiological responses to their highly dynamic arboreal habitat.     

Functional Analysis of the Intrathalline and Intracellular Chlorophyll Concentrations in the Lichen Family Umbilicariaceae

Intrathalline and intracellular chlorophyll concentrations togetherwith thallus structure were studied in 12 species of the lichenfamily Umbilicariaceae in order to explore pigment limitationsfor light absorption and for maximum rates of net photosynthesis(A_max). Species from high light environments tended to haverelatively low chlorophyll concentrations and vice versa. Chlorophyllconcentration on a surface area basis of all species studiedwas sufficient to absorb 80–99% of the incident light,which led us to believe that chlorophyll concentration was nota limiting factor for A_max. A_maxof Umbilicariaceae was smallerthan A_maxof leaves of vascular plants, both on a dry weightand on a surface area basis. These differences slightly decreasedbut did not disappear when referred to the chlorophyll concentrationof the photosynthetic tissues, but reference units had a confoundingeffect in the comparisons. Differences in A_maxon a dry weightbasis between Umbilicariaceae and vascular plants disappearedwhen comparisons were made with the whole plant and not simplywith the leaf. We found a significant, inverse relationshipbetween amount of alga and chlorophyll concentration withinthe algal cells in the lichens studied. This was interpretedas a mechanism to avoid pigment self-shading within the algallayer, distributing more efficiently the photosynthetic pigmentsover a given surface area, and is the first indication of sucha mechanism in lichens. It is suggested that thallus structureand physiology have a larger influence on the observed low A_maxofthe lichens studied than chlorophyll content. Algal cells; chlorophyll concentration; lichens; light harvesting; net photosynthesis; stereology; thallus structure; Umbilicariaceae     

A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance

Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M _A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F _L) modify light harvesting significantly. F _L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F _L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M _A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M _A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M _A, reductions in F _L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.     

Niche Differentiation in Tank and Atmospheric Epiphytic Bromeliads of a Seasonally Dry Forest

Factors influencing the niche differentiation of epiphytes have been determined for the epiphytic bromeliads that coexist in the seasonally dry forest of Chamela, Mexico. Over 40 percent of the bromeliad epiphytes were distributed in only 5 percent of the trees. The occurrence of compound leaves in host trees was highly correlated with abundance of epiphytes, as these allow scattered light to penetrate throughout the canopy. The effect of leaf type overrides the effect of bark type, the main factor determining seedling establishment in moist forests. Eight species had the atmospheric life form, while only two species had tanks, formed by overlapping leaf bases and associated to a lower drought tolerance. Distribution in the canopy is counter to that observed in moist forests, since tank species occur in the upper canopy. Tank life forms showed most annual carbon gain during the rainy season, when the newly leafed out trees provide shade to the lower canopy. Atmospheric species had photosynthetic activity for longer into the dry period, possibly supported by dew and fog events. Leaf angles, orientation, trichome, and stomata densities are discussed in relation to water and light use among the species with contrasting ecological strategies.     

The geometry of light interception by shoots of Heteromeles arbutifolia: morphological and physiological consequences for individual leaves

The influence of leaf orientation and position within shoots on individual leaf light environments, carbon gain, and susceptibility to photoinhibition was studied in the California chaparral shrub Heteromeles arbutifolia with measurements of gas exchange and chlorophyll fluorescence, and by application of a three-dimensional canopy architecture model. Simulations of light absorption and photosynthesis revealed a complex pattern of leaf light environments and resulting leaf carbon gain within the shoots. Upper, south-facing leaves were potentially the most productive because they intercepted greater daily photon flux density (PFD) than leaves of any other orientation. North-facing leaves intercepted less PFD but of this, more was received on the abaxial surface because of the steep leaf angles. Leaves differed in their response to abaxial versus adaxial illumination depending on their orientation. While most had lower photosynthetic rates when illuminated on their abaxial as compared to adaxial surface, the photosynthetic rates of north-facing leaves were independent of the surface of illumination. Because of the increasing self-shading, there were strong decreases in absorbed PFD and daily carbon gain in the basipetal direction. Leaf nitrogen per unit mass also decreased in the basipetal direction but on a per unit area basis was nearly constant along the shoot. The decrease in leaf N per unit mass was accounted for by an increase in leaf mass per unit area (LMA) rather than by movement of N from older to younger leaves during shoot growth. The increased LMA of older lower leaves may have contributed directly to their lower photosynthetic capacities by increasing the limitations to diffusion of CO₂ within the leaf to the sites of carboxylation. There was no evidence for sun/shade acclimation along the shoot. Upper leaves and especially south-facing upper leaves had a potential risk for photoinhibition as demonstrated by the high PFDs received and the diurnal decreases in the fluorescence ratio F _v/F _m. Predawn F _v/F _m ratios remained high (>0.8) indicating that when in their normal orientations leaves sustained no photoinhibition. Reorientation of the leaves to horizontal induced a strong sustained decrease in F _v/F _m and CO₂ exchange that slowly recovered over the next 10–15?days. If leaves were also inverted so that the abaxial surface received the increased PFDs, then the reduction in F _v/F _m and CO₂ assimilation was much greater with no evidence for recovery. The heterogeneity of responses was due to a combination of differences between leaves of different orientation, differences between responses on their abaxial versus adaxial surfaces, and differences along the shoot due to leaf age and self-shading effects.     

Plant growth-form alters the relationship between foliar morphology and species shade-tolerance ranking in temperate woody taxa

Variation in leaf size (area per leaf) and leaf dry weight per area (LWA) in relation to species shade- and drought-tolerance, characterised by Ellenberg's light (ELD) and water demand (EWD) values, respectively, were examined in 60 temperate woody taxa at constant relative irradiance. LWA was independent of plant size, but leaf size increased with total plant height at constant ELD. Canopy position also affected leaf morphology: leaves from the upper crown third had higher LWA and were larger than leaves from the lower third. Leaf size and LWA were negatively correlated, and leaf size decreased and LWA increased with decreasing species shade-tolerance. Mean LWA was similar for trees and shrubs, but trees had larger leaves than shrubs. Furthermore, all relationships were altered by plant growth-form: none of the qualitative tendencies was significant for trees. This implies the considerably lower plasticity of foliar parameters in trees than those in shrubs. Accordingly, shade-tolerance of trees, having relatively constant leaf structure, may be most affected by the variability in biomass partitioning and crown geometry which influence foliage distribution and spacing and finally determine canopy light absorptance. Alteration of leaf form and investment pattern for construction of unit foliar surface area which change the efficiency of light interception per unit biomass investment in leaves, is a competitive strategy inherent to shrubs. EWD as well as wood anatomy did not control LWA and leaf size, though there was a trend of ring-porous tree species to be more shade-tolerant than diffuse-porous trees. Since ring-porous species are more vulnerable to cavitation than diffuse-porous species, they may be constrained to environments where irradiances and consequently evaporative demand is lower.