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.     

PHYSIOLOGICAL AND MORPHOLOGICAL MODIFICATIONS OF PLANTAGO MAJOR (PLANTAGINACEAE) IN RESPONSE TO LIGHT CONDITIONS

Physiological and morphological differences between Plantago major L. (Plantaginaceae) growing in full sunlight and shaded conditions were examined. Photosynthesis of isolated leaves was saturated by irradiance around 300 μE m^−-2 sec^−-1 and 170 μE m^−-2 sec^−-1, respectively. In contrast to previous studies of sun/shade leaf responses, initial slopes of curves from shaded plants are significantly less than those taken from full-sun plants. Within the 400–500 nm and 600–700 nm ranges, leaves 5.0 cm or longer are essentially opaque, transmitting less than 1.25% of incident light. Chlorophyll content per unit leaf area is nearly equivalent for leaves from plants growing under the two extremes in light levels. Morphometric comparisons indicate shaded plants bear fewer leaves, have less leaf overlap, lower total leaf area, and longer petioles than full-sun plants. Leaf elongation rates are lower and the duration between the emergence of successive leaves is longer in shaded plants. Computer analyses of both types of rosette morphology reveal shaded plants have an equal or greater capacity to intercept light than full-sun plants, principally because of the minimization of leaf overlap and the large variation in the deflection angles of leaves in shaded rosette morphologies. Simulations, calculated on the basis of light interception, and taking into account the transition between photosynthate-importing and -exporting leaves, predict relative growth rates for full-sun and shaded rosette morphologies that are in reasonable agreement with empirically determined leaf growth rates. However, the data indicate that significant physiological and morphological differences exist among leaves from a single rosette, and among developmentally comparable leaves from rosettes growing under different ambient light environments. Differences among leaves on a single plant must be accommodated in computerized techniques attempting to simulate light interception and its consequences on potential growth rates.     

Leaf Heteroblasty in Pseudopanax crassifolius: Functional Significance of Leaf Morphology and Anatomy

Pseudopanax crassifolius (A. Cunn.) C. Koch. is a strongly heteroblastictree indigenous to New Zealand. Changes in the morphology andanatomy of leaves are described for seedling, juvenile, transitionaland adult phases of development. Seedlings produce five leaftypes; all are relatively small, thin, anthocyanic, have a lowspecific weight, and are anatomically comparable to the leavesof many shade plants. Juvenile leaves are long, linear, deflexed,coriaceous and sharply-toothed. They have a high specific weight,a thick, ornamented cuticle, a multiseriate hypodermis composedof collenchyma, well developed palisade and many spongy mesophylllayers. Juvenile leaves are strong relative to adult leavesand to the leaves of other species. Leaf strength is providedby fibres associated with the midrib. Adult leaves are shorter,broader and less massive than juvenile leaves, and are orientedhorizontally. Transitional leaves are morphologically intermediatebetween juvenile and adult leaves. The anatomy of juvenile,transitional and adult leaves is similar. The heteroblasticseries may be associated with changes in leaf construction costs,light interception and heat dissipation.Copyright 1993, 1999Academic Press Lancewood, Pseudopanax crassifolius (A. Cunn.) C. Koch., leaf heteroblasty, anatomy, morphology, structural adaptations     

Morphological responses of Datura ferox L. seedlings to the presence of neighbours

Summary We studied the effects of density on the dynamics of seedling growth and canopy microclimate within experimental stands composed of Datura ferox L. seedlings grown in individual pots. Interception of photosynthetically active radiation (PAR) by seedlings was evaluated either indirectly, by measuring leaf area, proportion of leaf area shaded by neighbouring individuals and laminar orientation with respect to sunlight, or directly, by measuring PAR at individual leaves at their natural angle of display. An integrating cylinder, with a geometry approximating that of a stem, was used within the canopies to measure the red:far-red (R:FR) ratio of the light flux from all compass points parallel to the soil surface. Seedlings responded rapidly (i.e. 1–2 weeks) to increased density by producing longer internodes and partitioning more dry matter to stems relative to leaves. These responses were observed before either PAR interception of growth rate were reduced by the presence of neighbours. Conversely, morphogenetic adjustment was preceded by a consistent effect of plant density on the R:FR ratio of the light received by the integrating cylinder. Air and soil temperature were not affected by density in these experiments. Differences in wind velocity within the canopy associated with plant density were avoided by the experimental procedure. The results support the idea that the drop in R:FR ratio of the light flux parallel to the ground — e.g. reflected sunlight — is an early signal that allows rapid adjustment of plant form to changes in canopy structure.     

Improved growth and water use efficiency of cherry saplings under reduced light intensity

Cherry (Prunus avium L.) saplings were grown under natural sunlight (controls) or moderate shading (up to 30%, depending on the incident light intensity and the hour of the day). Reduced light intensity increased the dry mass of each of the plant components studied. Consequently, the total dry mass of shaded plants was significantly greater than that of controls at the end of the growing season. However, the diurnal trend in the level of photosynthesis (per unit of leaf area) of shaded plants was similar to the controls in August, but lower in September. As the growing season proceeded, reduced photosynthetic rates, thinner mesophyll and larger specific leaf area in the shaded plants indicated that leaf development had adapted to shaded conditions throughout the growing season. It is suggested that increased growth of shaded plants was caused by a higher initial relative growth rate and a greater whole-plant photosynthesis. Shading consistently reduced transpiration over the season, therefore improving water use efficiency of shaded leaves. Our results suggest that a moderate reduction in light intensity can be a useful method for improving growth and saving water in hot and dry environments.     

Drought and changes in leaf orientation for two California chaparral shrubs: Ceanothus megacarpus and Ceanothus crassifolius

Summary Predawn xylem pressure potentials were measured on two California chaparral shrubs, Ceanothus megacarpus and Ceanothus crassifolius, throughout the winter and spring growing season and into the summer drought. On the days xylem pressure potentials were measured, leaf orientation measurements were made on a population of marked leaves from the same shrubs. Predawn xylem pressure potentials decreased from -0.1 MPa in both species to -7.8 and -6.6 MPa in C. megacarpus and C. crassifolius, respectively, between May and August, 1981. Leaf inclinations became more vertical during this period with the greatest change observed in C. crassifolius. This change in leaf inclination was reversible, and, in the late winter and early spring, one year old leaves became more horizontal. Leaf azimuths were random and did not change seasonally. Simulations of solar radiation interception indicated that the increase in leaf inclination associated with summer drought reduced the absorption of solar radiation in August by 6% for C. megacarpus and 20% for C. crassifolius. Standard leaf energy budget calculations suggest that steep leaf inclinations would result in slightly lower leaf temperatures and transpiration rates under summer conditions.     

LEAF ORIENTATION,RADIATION INTERCEPTION,AND NOCTURNAL ACIDITY INCREASES BY THE CAM PLANT AGAVE DESERTI (AGAVACEAE)

Both field measurements and a computer model were used to study the interception of photosynthetically active radiation (PAR) by Agave deserti (Engelm.), a desert CAM plant with a basal rosette of massive opaque leaves. PAR interception was determined in the winter and the summer for upper and lower leaf surfaces on a plant with about 60 leaves. Total daily PAR on the leaf surfaces was approximately 10 mol m^-2 for a winter day and 20 mol m^-2 for a summer day. For a PAR of 15 mol m^-2, the nocturnal increase in acidity was about 0.6 mol m^-2 for both leaf surfaces and various leaf orientations, except for the oldest most horizontal leaves where the increase was less than half as large. The acidity increase measured in the field was 90% saturated at 25 mol m^-2. Thus, daytime PAR in the desert is often limiting for the nocturnal acidity increase, especially for the lower leaf surfaces. Simulated tilting of the plant by 55° so that the vertical axis pointed to the sun at solar noon on a winter day increased the PAR incident on the upper surfaces of the leaves, but did not affect the total nocturnal increase in acidity by the whole plant. Although simulated removal of alternate leaves increased the PAR per unit leaf area for the remaining leaves, it reduced the total increase in nocturnal acidity of the whole plant by 31%. PAR interception by plants on slopes facing steeply north, east, or west was substantially reduced compared to the horizontal. Thus, the model proved to be quite useful for quantifying the relation between leaf orientation, PAR interception, and nocturnal increases in acidity by A. deserti, and it indicated that the lower frequency of plants on north- compared to south-facing slopes was due to PAR limitations.     

Leaf display and photosynthesis of tree seedlings in a cool-temperate deciduous broadleaf forest understorey

To examine a possible convergence in leaf photosynthetic characteristics and leaf display responses to light environment in seedlings of three canopy and two shrub tree species in understorey of cool-temperate deciduous broadleaf forest, relationships between light environment, leaf orientation and leaf light-photosynthetic response were measured. Light capture of the seedlings (17-24 individuals with 2-12 leaves for each species) was assessed with a three dimensional geometric modeling program Y-plant. Leaf photosynthetic characteristics of the five species were found to have acclimated to the understorey light environment, i.e., low light compensation point and high apparent quantum yield. In addition, light-saturated photosynthetic rates were higher in seedlings inhabiting microsites with higher light availability. Efficiencies of light capture and carbon gain of the leaf display were evaluated by simulating the directionalities of light capture and daily photosynthesis for each seedling using hemispherical canopy photography. The results showed that most of the seedlings orientated their leaves in a way to increase the daily photosynthesis during the direct light periods (sunflecks) rather than maximize daily photosynthesis by diffuse light. Simulations also showed that daily photosynthesis would increase only 10% of that on actual leaf display when the leaves orientated to maximize the diffuse light interception. Simulations in which leaf orientations were varied showed that when the leaf display fully maximized direct light interception, the time that leaves were exposed to excessive photon flux density of >800 mumol photons m(-2) s(-1) were doubled. The understorey seedlings studied responded to the given light environments in a way to maximize the efficiency of acquisition and use of light during their short (approximately 3 month) seasonal growth period.     

Screening a cucumber crop during leaf area development reduces yield

In order to reduce heat energy consumption in greenhouse cucumber production, (transparent) screens may be used also during the day, particularly in the early growth phase when high temperatures are required to achieve rapid leaf area development. However, energy savings must be optimised against light reduction‐induced yield loss. For this reason, two experiments were conducted to quantify the effect on photosynthesis and growth of screening cucumber plants during their early growth phase, and on yield in the following generative phase. Screening with different light transmission coefficients was simulated using shading nets. Shading the plants during the first 5 weeks under Central European winter conditions reduced the leaf area by 0.40% per 1% reduction in photosynthetic active radiation (PAR). Moreover, potential leaf net photosynthesis decreased by 0.46% per 1% PAR reduction. A major impact was that the leaf dry matter content, leaf starch content and leaf sugar content of shaded plants diminished significantly. In the course of the following 2 weeks under full light, the leaf photosynthesis of the plants previously shaded recovered fully and the leaf area index rose to 3.3 m² m^?2, considered sufficient for optimal crop photosynthesis. The yield from plants previously shaded diminished slightly as early as from the first harvest week on. These yield losses increased further over the next few weeks, measuring approximately 0.8 kg m^?2 per 1 mol m^?2 day^?1 PAR reduction in the early growth phase. The effect of PAR on plant growth was proportional when relating the PAR integral over the entire experimental period to the total yield and to the total dry matter production, respectively.     

Photosynthetically active radiation and carbon gain drives the southern orientation of Myrtillocactus geometrizans fruits

• The equatorial orientation of reproductive structures is known in some columnar cacti from extratropical deserts. It has been hypothesised that photosynthetically active radiation (PAR) interception is the main reason for this orientation, because of its key effect on nocturnal CO₂ uptake. However, there are no studies addressing both the effect of PAR and its consequence, carbon gain, on fruit orientation. Accordingly, we tested whether PAR and carbon gain could explain the southern fruit orientation of Myrtillocactus geometrizans, an inter‐tropical columnar cactus.
• We studied three populations of M. geometrizans in Mexico. For each population, azimuth of fruits, total daily PAR, nocturnal acid accumulation (NAA) and fruit production were measured. The relationships between rib orientation and number of fruits, as well as total daily PAR, were evaluated using periodic regressions. The effect of total daily PAR and NAA on number of fruits was assessed using generalised linear models.
• During spring, mean fruit orientation had a south azimuth for three populations. Likewise, rib orientation had a significant effect on fruit production, with the south‐facing ribs having the maximum number of fruits. Total daily PAR was highest in the south‐facing ribs, at least for those in the northern and central populations. Furthermore, during spring, there was a significant positive effect of total daily PAR and NAA on fruit production.
• Our results provide strong evidence that the higher carbon gain in equatorial ribs, through a highest interception of PAR, would be the responsible factor for equatorial orientation of fruits in an inter‐tropical columnar cactus.

     

Canopy structure,light microclimate and leaf gas exchange of Quercus coccifera L. in a Portuguese macchia: measurements in different canopy layers and simulations with a canopy model

Summary The structural characteristics of a diverse array of Quercus coccifera canopies were assessed and related to measured and computed light attenuation, proportion of sunlit foliage, foliage temperatures, and photosynthesis and diffusive conductance behavior in different canopy layers. A canopy model incorporating all components of shortwave and longwave radiation, and the energy balance, conductance, and CO₂ and H₂O exchanges of all leaf layers was developed and compared with measurements of microclimate and gas exchange in canopies in four seasons of the year. In the denser canopies with a leaf area index (LAI) greater than 5, there is little sunlit foliage and the diffuse radiation (400–700 nm) is attenuated to 5% or less of the global radiation (400–700 nm) incident on the top of the canopy. Foliage of this species is nonrandomly distributed with respect to azimuth angle, and within each canopy layer, foliage azimuth and inclination angles are correlated. A detailed version of the model which computed radiation interception and photosynthetic light harvesting according to these nonrandom distributions indicated little difference in whole-canopy gas exchange from calculations of the normal model, which assumes random azimuth orientation. The contributions of different leaf layers to canopy gas exchange are not only a function of the canopy microclimate, but also the degree to which leaves in the lower layers of the canopy exhibit more shade-leaf characteristics, such as low photosynthetic and respiratory capacity and maximal conductance. On cloudless days, the majority of the foliage in a canopy of 5.4 LAI is shaded —70%–90% depending on the time of year. Yet, the shaded foliage under these conditions is calculated to contribute only about one-third of the canopy carbon gain. This contribution is about the same as that of the upper 13% of the canopy foliage. Computed annual whole-canopy carbon gain and water use are, respectively, 60% and 100% greater for a canopy of 5 LAI than for one of 2 LAI. Canopy water-use efficiency is correspondingly less for the canopy of 5 LAI than for that of 2 LAI, but most of this difference is apparent during the cool months of the year, when moisture is more abundant.     

Foliole movement and canopy architecture of Larrea tridentata (DC.) Cov. in Mexican deserts

Summary The creosote bush (Larrea tridentata) is a common desert perennial with bifoliate, amphistomatic, divaricate leaves. The leaves can vertically close their folioles and vary their profile with respect to direct solar radiation. Field data from different Mexican deserts showed a significant correlation between foliole aperture and mean foliole inclination: in plants in which folioles were more open, the foliole surfaces were less vertical. In a series of field experiments in the Chihuahuan Desert, foliole aperture varied significantly with the water-status of the plant and the hour of the day. In moist plants, folioles opened in the early morning and closed in the afternoon. Water-stressed plants showed significantly lower foliole apertures. A simulation of the light interception patterns of the plants showed that foliole closure in water-stressed individuals reduces direct radiation interception by around 24%. Most (64%) of the reduction in interception was due to the vertical inclination of the photosynthetic surfaces induced by foliole closure in the water-stressed plants. The rest (36%) of the reduction in interception was due to differential self-shading between foliole pairs, which was higher in the closed folioles of the water-stressed plants, but operated more towards the early hours of the day.     

Natural selection on ecophysiological traits of a fern species in a temperate rainforest

Unlike other species of the genus Blechnum, the fern Blechnum chilense occurs in a wide range of habitats in Chilean temperate rainforest, from shaded forest understories to abandoned clearings and large gaps. We asked if contrasting light environments can exert differential selection on ecophysiological traits of B. chilense. We measured phenotypic selection on functional traits related to carbon gain: photosynthetic capacity (A _max), dark respiration rate (R _d), water use efficiency (WUE), leaf size and leaf thickness in populations growing in gaps and understorey environments. We assessed survival until reproductive stage and fecundity (sporangia production) as fitness components. In order to determine the potential evolutionary response of traits under selection, we estimated the genetic variation of these traits from clonally propagated individuals in common garden experiments. In gaps, survival of B. chilense was positively correlated with WUE and negatively correlated with leaf size. In contrast, survival in shaded understories was positively correlated with leaf size. We found positive directional fecundity selection on WUE in gaps population. In understories, ferns of lower R _d and greater leaf size showed greater fecundity. Thus, whereas control of water loss was optimized in gaps, light capture and net carbon balance were optimized in shaded understories. We found a significant genetic component of variation in WUE, R _d and leaf size. This study shows the potential for evolutionary responses to heterogeneous light environments in functional traits of B. chilense, a unique fern species able to occupy a broad successional niche in Chilean temperate rainforest.     

LEAF DISPLAY,CANOPY STRUCTURE,AND LIGHT INTERCEPTION OF TWO UNDERSTORY PALM SPECIES

The implications of leaf size, leaf display, and crown size for whole-plant light interception were investigated in Geonoma cuneata and Asterogyne martiana, two understory palm species native to Central American rain forests. Adults of A. martiana had longer leaves, more leaves per plant, and greater total leaf area than G. cuneata adults. Geometric measurements within whole crowns were used to calculate light interception efficiency, a leaf-based measure of the proportion of total incident light that is intercepted by a crown. Light interception efficiency was higher in adult G. cuneata than in adult A. martiana; seedlings of the two species did not differ significantly in light interception efficiency. Decreased efficiency of adult A. martiana crowns was largely due to an increased proportion of pendent leaves. Compared to G. cuneata, adults of A. martiana had greater light interception capacity (the product of light interception efficiency and total leaf area), but they also had higher biomass costs of light interception. Lower biomass costs of light interception in adult G. cuneata enable this species to exploit successfully the most deeply shaded microsites in the rain forest understory.     

Daylight spectra (400–740 nm) beneath sunny,blue skies in Tasmania,and the effect of a forest canopy

Abstract The photosynthetically active radiation (PAR) incident on a horizontal surface at an open mountain site is positively correlated with solar altitude for sunny, blue sky conditions. The proportion of red light in PAR decreases with increasing solar altitude, while that of blue increases. These results are consistent with the wavelength dependency of Rayleigh and Mie scattering. The ratio of near infrared radiation to PAR decreases with increasing solar altitude towards solar noon and with decreasing solar altitude towards sunset. Thus surface reflection seems to be an important part of the light climate. The relative transmission of daylight through a forest canopy to a horizontal surface is not correlated with solar altitude for sunny, blue sky conditions at a mountain site. The amount of diffuse daylight is negatively correlated with per cent canopy interception, and the amount of direct sunlight is negatively correlated with per cent solar track interception. Daylength is negatively correlated with both canopy and solar track interceptions. The proportion of red light in PAR increases with increasing solar altitude, while that of blue decreases. These results are opposite those for the open site and are due to the spatial patterns of canopy obstruction of the sky vault, and of the spectral quality of daylight across the sky. The ratio of near infrared radiation to PAR in shadelight increases with increasing canopy interception due to the selective scattering properties of the canopy. The ratio for shadelight is positively correlated with the ratio for sunflecks.     

Plant competition for light analyzed with a multispecies canopy model

Summary Competition for light among species in a mixed canopy can be assessed quantitatively by a simulation model which evaluates the importance of different morphological and photosynthetic characteristics of each species. A model was developed that simulates how the foliage of all species attenuate radiation in the canopy and how much radiation is received by foliage of each species. The model can account for different kinds of foliage (leaf blades, stems, etc.) for each species. The photosynthesis and transpiration for sunlit and shaded foliage of each species is also computed for different layers in the canopy. The model is an extension of previously described single-species canopy photosynthesis simulation models. Model predictions of the fraction of foliage sunlit and interception of light by sunlit and shaded foliage for monoculture and mixed canopies of wheat (Triticum aestivum) and wild oat (Avena fatua) in the field compared very well with measured values. The model was used to calculate light interception and canopy photosynthesis for both species of wheat/wild oat mixtures grown under normal solar and enhanced ultraviolet-B (290–320 nm) radiation (UV-B) in a glasshouse experiment with no root competition. In these experiments, measurements showed that the mixtures receiving enhanced UV-B radiation had a greater proportion of the total foliage area composed of wheat compared to mixtures in the control treatments. The difference in species foliage area and its position in the canopy resulted in a calculated increase in the portion of total canopy radiation interception and photosynthesis by wheat. This, in turn, is consistent with greater canopy biomass of wheat reported in canopies irradiated with supplemental UV-B.     

THE ROLE OF PHYLLOTACTIC PATTERN AS A “DEVELOPMENTAL CONSTRAINT” ON THE INTERCEPTION OF LIGHT BY LEAF SURFACES

Computer simulations of model plants are used to assess the influence of leaf shape, size, and pattern of arrangement (= phyllotaxy) on the direct solar radiation intercepted by leaf surfaces. Changes in phyllotaxy significantly influence light interception (and, by inference, net assimilation rate) for rosette growth habits. However, changes in leaf shape and orientation and in stem length can compensate for the negative effects of leaf overlap produced by phyllotactic patterns. Phyllotaxy is viewed as a developmental limiting factor in photobiology that may necessitate compensatory changes in other morphological features not directly controlled by patterns of leaf initiation. This distinguishes it from functioning as a “developmental constraint” sensu stricto and may provide a paradigm for other features in plant evolution.     

Effects of light quality and quantity on growth of the clonal plant Eichhornia crassipes

Summary Plant canopy shade reduces photosynthetic photon flux density (PPFD) and ratio of red to far-red light (z). Both effects can cause plants to increase potential for light acquisition through vertical growth and leaf area expansion. Clonal plants such as Eichhornia crassipes might alternatively increase light interception via horizontal growth of stolons or rhizomes and placement of new ramets in less shaded microsites. Effect of simulated canopy shade and component effects of PPFD and z were tested by filtering or adding light uniformly, to a whole group of connected ramets, or locally, to individual ramets within a group. In uniform treatments, low PPFD reduced total growth but low z did not. Low PPFD and low z independently reduced stolon and ramet production and caused etiolation of petioles; effect of low PPFD plus low z on ramet production was greater than that of either factor alone. Lateral clonal growth thus did not seem to be a response to uniform shading; instead, uniformly low PPFD or low z increased partitioning to established ramets. Low z changed partitioning without changing total growth. In local treatments, reduction of growth of individual ramets due to low PPFD and inhibition of new ramet production attributable to spectral composition of light were mitigated when connected ramets were unshaded; plants may respond differently to patchy than to uniform shade.     

Herbivory by leaf miners in response to experimental shading of a native crucifer

Summary We tested the hypothesis that light intensity was the direct, proximal mechanism causing significantly higher vulnerability of Bittercress (Cardamine cordifolia A. Gray) clones in the sun to herbivory by a leaf-mining fly (Scaptomyza nigrita Wheeler). Clones in the sun were experimentally shaded. Plant performance and losses to leaf miners were compared to controls in the sun and natural willow shade. Leaf-mining damage was significantly higher on artificially-shaded plants (P<0.01), opposite of our expectation. Shading sun plants shifted their growth pattern toward that of naturally-shaded plants. No significant differences were detected in leaf water status or glucosinolate concentrations, eliminating water stress and variation in defensive posture for mediating the between habitat differences in levels of herbivory. Although soluble sugars varied significantly, they were higher in sun than either shade treatment. Total and free amino nitrogen concentrations were highest in the artificially-shaded plants and lowest in naturally-occurring sun plants. Adult flies were more abundant on sun and on artificially-shaded plants than on naturally-shaded plants. Thus, relative abundance of ovipositing flies in the sun-exposed area, combined with the higher nitrogen availability in artificially-shaded plants, form the most plausible hypothesis for factors mediating the experimentally documented pattern of herbivory.     

Modeling the light interception and carbon gain of individual fluttering aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis> Michx) leaves

Poplars (Populus spp.) have a particular petiole morphology that enhances leaf flutter even in light winds. Previous studies have shown that this trait enhances whole canopy carbon gain through changes in the temporal dynamics and spatial distribution of light in the lower canopy. However, less is known about the effects of flutter for leaves at the top of the canopy ("sun leaves"). A computer simulation model was developed that uses latitude, time of day, day of year, azimuth and a slope component, which was varied at a 3 Hz frequency over an arc of rotation to create the flutter motion, and generate data on light interception for both surfaces of a fixed or fluttering leaf. The light files generated (10 Hz) were input into a dynamic model of photosynthesis to estimate the carbon gain for both fluttering and fixed leaves. As compared to leaves fixed at various angles and azimuths, fluttering leaves had a more uniform light interception. Depending on their angle and azimuth, fixed leaves may not always be intercepting high photon flux density (PFD) even when exposed to full sun. Leaf flutter continuously randomizes leaf angles creating uniform PFD inputs for photosynthetic reactions regardless of variation in leaf orientation and solar position. These effects on light interception could have positive impacts on carbon gain for leaves at the top of the canopy.