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Comparative locomotor ecology of gibbons and macaques: selection of canopy elements for crossing gaps.

To examine functional questions of arboreal locomotor ecology, the selection of canopy elements by Bornean agile gibbons (Hylobates agilis) and long-tailed macaques (Macaca fascicularis) was contrasted, and related to locomotor behaviors. The two species, and in some cases, the macaque sexes, varied in their use of most structural elements. Although both species traveled most frequently in the main canopy layer (macaques: 56%, gibbons: 48%), the gibbons strongly preferred the emergent canopy layer and traveled higher than the macaques (31 vs. 23 m above ground) in larger trees (48 vs. 26 cm dbh). Macaques preferred to cross narrower gaps (50% were in the class 0.1-0.5 m wide) than gibbons (42% were 1.6-3.0 m wide), consistent with the maximum gap width each crossed (3.5 m for macaques, 9 m for gibbons). Macaques could cross only 12% of the gaps encountered in the main canopy, and < 5% of the gaps in each of the other four layers. In contrast, all layers appear relatively continuous for gibbons. Specialized locomotor modes were used disproportionately at the beginning and end of travel segments, further indicating that behavior was organized around gap crossings. A model is defined, the Perceived Continuity Index (PCI), which predicts the relative use of canopy strata for each species, based on the percentage of gaps a species can cross, the frequency of gaps, and median length of continuous canopy structure in each canopy layer. The results support the hypothesis that locomotor behaviors, and strategies of selecting canopy strata for travel, are strongly constrained by wide gaps between trees and are ultimately based on selection for efficient direct line travel between distant points.

Animals↗

Effects of air velocity on photosynthesis of plant canopies under elevated CO2 levels in a plant culture system.

To obtain basic data for adequate air circulation for promoting plant growth in closed plant production modules in bioregenerative life support systems in space, effects of air velocities ranging from 0.1 to 0.8 m s-1 on photosynthesis in tomato seedlings canopies were investigated under atmospheric CO2 concentrations of 0.4 and 0.8 mmol mol-1. The canopy of tomato seedlings on a plug tray (0.4 x 0.4 m2) was set in a wind-tunnel-type chamber (0.6 x 0.4 x 0.3 m3) installed in a semi-closed-type assimilation chamber (0.9 x 0.5 x 0.4 m3). The net photosynthetic rate in the plant canopy was determined with the differences in CO2 concentrations between the inlet and outlet of the assimilation chamber multiplied by the volumetric air exchange rate of the chamber. Photosynthetic photon flux (PPF) on the plant canopy was kept at 0.25 mmol m-2 s-1, air temperature at 23 degrees C and relative humidity at 55%. The leaf area indices (LAIs) of the plant canopies were 0.6-2.5 and plant heights were 0.05-0.2 m. The net photosynthetic rate of the plant canopy increased with increasing air velocities inside plant canopies and saturated at 0.2 m s-1. The net photosynthetic rate at the air velocity of 0.4 m s-1 was 1.3 times that at 0.1 m s-1 under CO2 concentrations of 0.4 and 0.8 mmol mol-1. The net photosynthetic rate under CO2 concentrations of 0.8 mmol mol-1 was 1.2 times that under 0.4 mmol mol-1 at the air velocity ranging from 0.1 to 0.8 m s-1. The results confirmed the importance of controlling air movement for enhancing the canopy photosynthesis under an elevated CO2 level as well as under a normal CO2 level in the closed plant production modules.

Air Conditioning↗

Canopy development, CO(2) exchange and carbon balance of a modeled agroforestry tree.

We developed a whole-canopy CO(2) exchange simulation model to study effects of pruning on the carbon balance of trees. Model inputs include global short-wave radiation, photosynthetic photon flux density (PFD), air temperature, time series of the development of canopy diameter, height and total leaf area during the simulation period and local geographical and atmospheric parameters. Canopy structure is derived stochastically from the time series of canopy development and growth functions of individual phytoelements. The PFD incident on a phytoelement is computed from the average gap frequency of the canopy and the binary random probability of sunflecks on the phytoelement. Instantaneous CO(2) assimilation rate of each phytoelement is computed from PFD and phytoelement age. Assimilation rates are integrated over space and time to estimate whole-canopy CO(2) assimilation. The model was used to study carbon balance in five sources of the leguminous agroforestry tree Erythrina poeppigiana (Walpers) O.F. Cook during two 6-month pruning intervals. The canopy description appeared to be realistic. According to the simulations, cumulative assimilation did not provide enough carbon for tree growth until two months after pruning, indicating dependence of tree growth on reserve carbohydrates. The two most productive sources, which had the most open canopies, were the most dependent on reserve carbohydrates after pruning.

Journal Article↗

Vertical gradients in photosynthetic light response within an old-growth Douglas-fir and western hemlock canopy.

We examined needle-level light response of photosynthesis across a vertical light gradient within 45-55-m-tall western hemlock (Tsuga heterophylla (Raf.) Sarg.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees growing in a 400-500-year-old mixed species stand. We determined: (1) whether light-saturated photosynthetic rates, light compensation points, and respiration rates varied from the upper to the lower canopy, and (2) if light-saturated photosynthetic rates, light compensation points, and respiration rates varied between Douglas-fir and western hemlock. Over a 25-m gradient from the canopy top to the lower canopy, mean light-saturated photosynthetic rates, light compensation points, and respiration rates declined in overstory Douglas-fir and western hemlock needles, paralleling a 65% decline in the mean daily photosynthetic photon flux density (PPFD). At the canopy top, increasing light-saturated photosynthetic rates relative to lower canopy needles increased carbon uptake at high PPFD. In the lower canopy, reduced respiration rates relative to upper canopy needles increased carbon uptake at low PPFD by reducing the light compensation point. At all canopy positions, western hemlock had lower mean light-saturated photosynthetic rates, light compensation points and respiration rates than Douglas-fir. As a result, western hemlock had higher net photosynthetic rates at low PPFD, but lower net photosynthetic rates at high PPFD compared with Douglas-fir.

Journal Article↗

Leaf area distribution and radiative transfer in open-canopy forests: implications for mass and energy exchange.

Leaf area and its spatial distribution are key canopy parameters needed to model the radiation regime within a forest and to compute the mass and energy exchange between a forest and the atmosphere. A much larger proportion of available net radiation is received at the forest floor in open-canopy forests than in closed-canopy forests. The proportion of ecosystem water vapor exchange (lambda E) and sensible heat exchange from the forest floor is therefore expected to be larger in open-canopy forests than in closed-canopy forests. We used a combination of optical and canopy geometry measurements, and robust one- and three-dimensional models to evaluate the influence of canopy architecture and radiative transfer on estimates of carbon, water and energy exchange of a ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forest. Three-dimensional model simulations showed that the average probability of diffuse and direct radiation transmittance to the forest floor was greater than if a random distribution of foliage had been assumed. Direct and diffuse radiation transmittance to the forest floor was 28 and 39%, respectively, in the three-dimensional model simulations versus 23 and 31%, respectively, in the one-dimensional model simulations. The assumption of randomly distributed foliage versus inclusion of clumping factors in a one-dimensional, multi-layer biosphere-atmosphere gas exchange model (CANVEG) had the greatest effect on simulated annual net ecosystem exchange (NEE) and soil evaporation. Assuming random distribution, NEE was 41% lower, net photosynthesis 3% lower, total lambda E 10% lower, and soil evaporation 40% lower. The same comparisons at LAI 5 showed a similar effect on annual NEE estimates (37%) and lambda E (12%), but a much larger effect on net photosynthesis (20%), suggesting that, at low LAI, canopies are mostly sunlit, so that redistribution of light has little effect on net photosynthesis, whereas the effect on net photosynthesis is much greater at high LAIs.

Biomass↗

Age-related effects on leaf area/sapwood area relationships, canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge, Germany.

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.

Germany↗

Beetle assemblages from an Australian tropical rainforest show that the canopy and the ground strata contribute equally to biodiversity.

There remains great uncertainty about how much tropical forest canopies contribute to global species richness estimates and the relative specialization of insect species to vertical zones. To investigate these issues, we conducted a four-year sampling program in lowland tropical rainforest in North Queensland, Australia. Beetles were sampled using a trap that combines Malaise and flight interception trap (FIT) functions. Pairs of this trap, one on the ground and a second suspended 15-20 m above in the canopy were located at five sites, spaced 50 m or more apart. These traps produced 29986 beetles of 1473 species and 77 families. There were similar numbers of individuals (canopy 14473; ground 15513) and species (canopy 1158; ground 895) in each stratum, but significantly more rare species in the canopy (canopy 509; ground 283). Seventy two percent of the species (excluding rare species) were found in both strata. Using IndVal, we found 24 and 27% of the abundant species (n>or=20 individuals) to be specialized to the canopy and the ground strata, respectively, and equivalent analyses at the family level showed figures of 30 and 22%, respectively. These results show that the canopy and the ground strata both provide important contributions to rainforest biodiversity.

Animals↗

Modelling the effect of diffuse light on canopy photosynthesis in controlled environments.

A layered canopy model was used to analyze the effects of diffuse light on canopy gross photosynthesis in controlled environment plant growth chambers, where, in contrast to the field, highly diffuse light can occur at high irradiance. The model suggests that high diffuse light fractions (approximately 0.7) and irradiance (1400 micromoles m-2 s-1) may enhance crop life-cycle canopy gross photosynthesis for hydroponic wheat by about 20% compared to direct light at the same irradiance. Our simulations suggest that high accuracy is not needed in specifying diffuse light fractions in chambers between approximately 0.7 and 1, because simulated photosynthesis for closed canopies plateau in this range. We also examined the effect of leaf angle distribution on canopy photosynthesis under growth chamber conditions, as these distributions determine canopy extinction coefficients for direct and diffuse light. We show that the spherical leaf angle distribution is not suitable for modeling photosynthesis of planophile canopies (e.g., soybean and peanut) in growth chambers. Also, the absorption of the light reflected from the surface below the canopy should generally be included in model simulations, as the corresponding albedo values in the photosynthetically active range may be quite high in growth chambers (e.g., approximately 0.5). In addition to the modeling implications, our results suggest that diffuse light conditions should be considered when drawing conclusions from experiments in controlled environments.

Computer Simulation↗

Effects of light intensity and air velocity on air temperature, water vapor pressure, and CO2 concentration inside a plant canopy under an artificial lighting condition.

In order to characterize environmental variables inside a plant canopy under artificial lighting in the CELSS, we investigated the effects of light intensity and air velocity on air temperature, water vapor pressure, and CO2 concentration inside a plant canopy. Under a PPF of 500 micromoles m-2 s-1, air temperature was 2-3 degrees C higher, water vapor pressure was 0.6 kPa higher, and CO2 concentration was 25-35 micromoles mol-1 lower at heights ranging from 0 to 30 mm below the canopy than at a height 60 mm above the canopy. Increasing the PPF increased air temperature and water vapor pressure and decreased CO2 concentration inside the canopy. The air temperature was lower and the CO2 concentration was higher inside the canopy at an air velocity of 0.3 m s-1 than at an air velocity of 0.1 m s-1. The environmental variables inside the canopy under a high light intensity were characterized by higher air temperature, higher vapor pressure, and lower CO2 concentration than those outside the canopy.

Air Conditioning↗

Forest Canopy Stratification-Is It Useful?

It has long been recognized that the forest canopy has a complex structure that is significant for environmental interactions, regeneration, growth, and biotic habitat. Not only is the structure variously complex, but also there are many ways to conceptualize that complexity. Yet the persistent theme when considering the structure of canopies continues to be that of stratification: whether structural units are arranged in layers above the ground. We examined the use of the terms "stratification," "layering," and others in connection with canopy structure and found they had various meanings (often only implied) that were difficult to reconcile and to measure. We applied the definitions to the structure of a single, well-studied canopy located in Virginia, U.S.A., and found they failed to define consistently and clearly the presence, number, or location of strata. Additionally, we found the concept had limitations related to scale dependence, point of reference, and spatial averaging. Thus, asserting that a forest is stratified or naming the number of layers generally provided no guide to its structure. We propose alternative ways of conceptualizing and studying the forest canopy that avoid most of the problems associated with stratification. Among these are direct measurement and mapping of structural and environmental variables that have clear potential connections with canopy functions and viewing the distribution of structures or environmental conditions within the canopy as ecological gradients.

canopy↗

Form-function analysis of the effect of canopy morphology on leaf self-shading in the seagrass Thalassia testudinum.

The variation in seagrass morphology and the magnitude of leaf self-shading within the canopy of Thalassia testudinum, were compared among nine sites in a fringing reef lagoon. We found a significant variation in the growth-form of T. testudinum reflected in a 5.4-fold variation in the attenuation coefficient (K (d)) within the canopy. The largest morphological variation was observed in shoot density. Leaf biomass, leaf area index (LAI), and shoot density were positively associated with canopy-K (d) and with the percentage of surface irradiance received by the top of the seagrass canopy (% Es). These results provide an explanation for the consistent pattern of depth reduction in seagrass leaf biomass and shoot density reported in the literature. Shoot density and shoot size are two descriptors of the growth-form of T. testudinum related to its clonal life-form. Shoot size was not significantly correlated with canopy-K (d), nevertheless, it showed a significant effect on the slope of the relationship between shoot density and canopy-K (d). According to this model, shoot size also contributes to light attenuation within the seagrass canopy by increasing the effect of shoot density. This form-function analysis suggests that light may have a relevant role in the regulation of the optimal plant balance between horizontal (variation in shoot density) and vertical (variation in shoot size) growth of seagrasses. Other environmental factors and interactions also need to be examined to fully understand the mechanistic bases of the morphological responses of seagrasses to the environment.

Biomass↗

Physiology, morphology, and ozone uptake of leaves of black cherry seedlings, saplings, and canopy trees.

Patterns of ozone uptake were related to physiological, morphological, and phenological characteristics of different-sized black cherry trees (Prunus serotina Ehrh.) at a site in central Pennsylvania. Calculated ozone uptake differed among open-grown seedlings, forest gap saplings, and canopy trees and between leaves in the upper and lower crown of saplings and canopy trees. On an instantaneous basis, seedling leaves had the greatest ozone uptake rates of all tree size classes due to greater stomatal conductance and higher concentrations of ozone in their local environment. A pattern of higher stomatal conductance of seedlings was consistent with higher incident photosynthetically-active radiation, stomatal density, and predawn xylem water potentials for seedlings relative to larger trees. However, seedlings displayed an indeterminate pattern of shoot growth, with the majority of their leaves produced after shoot growth had ceased for canopy and sapling trees. Full leaf expansion occurred by mid-June for sapling and canopy trees. Because many of their leaves were exposed to ozone for only part of the growing season, seedlings had a lower relative exposure over the course of the growing season, and subsequently lower cumulative uptake, of ozone than canopy trees and a level of uptake similar to upper canopy leaves of saplings. Visible injury symptoms were not always correlated with patterns in ozone uptake. Visible symptoms were more apparent on seedling leaves in concurrence with their high instantaneous uptake rates. However, visible injury was more prevalent on leaves in the lower versus upper crown of canopy trees and saplings, even though lower crown leaves had less ozone uptake. Lower crown leaves may be more sensitive to ozone per unit uptake than upper crown leaves because of their morphology. In addition, the lower net carbon uptake of lower crown leaves may limit repair and anti-oxidant defense processes.

Journal Article↗

Acclimation response of spring wheat in a free-air CO(2) enrichment (FACE) atmosphere with variable soil nitrogen regimes. 3. Canopy architecture and gas exchange.

The response of whole-canopy net CO(2) exchange rate (CER) and canopy architecture to CO(2) enrichment and N stress during 1996 and 1997 for open-field-grown wheat ecosystem (Triticum aestivum L. cv. Yecora Rojo) are described. Every Control (C) and FACE (F) CO(2) treatment (defined as ambient and ambient +200 mumol mol(-1), respectively) contained a Low- and High-N treatment. Low-N treatments constituted initial soil content amended with supplemental nitrogen applied at a rate of 70 kg N ha(-1) (1996) and 15 kg N ha(-1) (1997), whereas High-N treatments were supplemented with 350 kg N ha(-1) (1996 and 1997). Elevated CO(2) enhanced season-long carbon accumulation by 8% and 16% under Low-N and High-N, respectively. N-stress reduced season-long carbon accumulation 14% under ambient CO(2), but by as much as 22% under CO(2) enrichment. Averaging both years, green plant area index (GPAI) peaked approximately 76 days after planting at 7.13 for FH, 6.00 for CH, 3.89 for FL, and 3.89 for CL treatments. Leaf tip angle distribution (LTA) indicated that Low-N canopies were more erectophile than those of High-N canopies: 48 degrees for FH, 52 degrees for CH, and 58 degrees for both FL and CL treatments. Temporal trends in canopy greenness indicated a decrease in leaf chlorophyll content from the flag to flag-2 leaves of 25% for FH, 28% for CH, 17% for CL, and 33% for FL during 1997. These results indicate that significant modifications of canopy architecture occurs in response to both CO(2) and N-stress. Optimization of canopy architecture may serve as a mechanism to diminish CO(2) and N-stress effects on CER.

Journal Article↗

Effects of canopy gaps on the genetic structure of Camellia japonica saplings in a Japanese old-growth evergreen forest.

The genetic structure of Camellia japonica saplings was investigated in relation to canopy conditions in an old-growth evergreen forest in Tsushima, Japan. To elucidate effects of canopy gaps on genetic structure, a 1 ha study site was divided into 20 x 20 m quadrats, which were classified into a gap quadrats (GAP), closed canopy quadrats (CLS) and mixed quadrats. Five GAP quadrats and six CLS quadrats were analyzed separately. Isolation-by-distance was tested by examining the correlation between genetic distance and geographic distance. A significant positive correlation was detected for GAP quadrats, whilst that for CLS quadrats was significantly smaller and not significantly different from zero. On the other hand, an analysis using Moran's I spatial autocorrelation coefficients indicates that the genetic structure is weaker in GAP quadrats than in CLS quadrats in short distance classes. The values were significantly positive for both types of quadrat. These results, along with our field observations on flowering, suggest that canopy gaps affect the genetic structure of C. japonica saplings in two distinct ways. First, canopy gaps may promote flowering and mating in an isolation-by-distance manner within canopy gaps. Second, canopy gaps may promote seed production and resulting overlap in seed shadows may weaken fine-scale genetic structures.

Camellia↗

The slow reversibility of photosystem II thermal energy dissipation on transfer from high to low light may cause large losses in carbon gain by crop canopies: a theoretical analysis.

Regulated thermal dissipation of absorbed light energy within the photosystem II antenna system helps protect photosystem II from damage in excess light. This reversible photoprotective process decreases the maximum quantum yield of photosystem II (Fv)/Fm) and CO2 assimilation (phiCO2), and decreases the convexity of the non-rectangular hyperbola describing the response of leaf CO2 assimilation to photon flux (theta). At high light, a decrease in phiCO2 has minimal impact on carbon gain, while high thermal energy dissipation protects PSII against oxidative damage. Light in leaf canopies in the field is continually fluctuating and a finite period of time is required for recovery of phiCO2 and when light drops below excess levels. Low phiCO2) and can limit the rate of photosynthetic carbon assimilation on transfer to low light, an effect prolonged by low temperature. What is the cost of this delayed reversal of thermal energy dissipation and phiCO2 recovery to potential CO2 uptake by a canopy in the field? To address this question a reverse ray-tracing algorithm for predicting the light dynamics of 120 randomly selected individual points in a model canopy was used to describe the discontinuity and heterogeneity of light flux within the canopy. Because photoprotection is at the level of the cell, not the leaf, light was simulated for small points of 10(4) micro m rather than as an average for a leaf. The predicted light dynamics were combined with empirical equations simulating the dynamics of the light-dependent decrease and recovery of phiCO2 and and their effects on the integrated daily canopy carbon uptake (A'c). The simulation was for a model canopy of leaf area index 3 with random inclination and orientation of foliage, on a clear sky day (latitude 44 degrees N, 120th day of the year). The delay in recovery of photoprotection was predicted to decrease A'c by 17% at 30 degrees C and 32% at 10 degrees C for a chilling-susceptible species, and by 12.8% at 30 degrees C and 24% at 10 degrees C for a chilling-tolerant species. These predictions suggest that the selection, or engineering, of genotypes capable of more rapid recovery from the photoprotected state would substantially increase carbon uptake by crop canopies in the field.

Carbon↗

Canopy photosynthesis and respiration of kiwifruit (Actinidia deliciosa var. deliciosa) vines growing in the field.

Net CO(2) assimilation (A) for canopies of kiwifruit (Actinidia deliciosa var. deliciosa) vines enclosed in a whole-canopy cuvette was measured continuously for three periods of 15-20 days during late summer, near Hamilton, New Zealand (latitude 38.2 degrees S). Canopy A showed an asymptotic response to incident radiation (PAR), saturating at about 1300 micromol m(-2) s(-1) for one vine and about 800 micromol m(-2) s(-1) for two other vines. Radiation interception at low solar angles and low leaf area apparently limited the response of A to PAR. Radiation saturated rates of A were 25-30 micromol CO(2) m(-2) s(-1) for one vine, and 12-18 micromol CO(2) m(-2) s(-1) for two other vines. At any PAR, canopy A was often lower in the afternoon than in the morning. Canopy respiration averaged 8.9 micromol CO(2) m(-2) s(-1) at 12 degrees C, but increased only 24-34% over the range 7-17 degrees C. Net daily C gains for the whole canopy, calculated as the temporal integral of A, ranged from -0.8 g C m(-2) for a cloudy day (PAR </= 450 micromol m(-2) s(-1)) to 6.2 g C m(-2) for a day with intermittent cloud. Measurements of gas exchange for single leaves in a leaf cuvette were used to examine the basis of the whole-canopy responses to environmental variables. Partial stomatal closure due to water stress apparently contributed to the afternoon depressions in A, although non-stomatal limitations may also have been involved.

Journal Article↗

Shoot structure, light interception, and distribution of nitrogen in an Abies amabilis canopy.

We studied the effects of variation in shoot structure and needle morphology on the distributions of light and nitrogen within a Pacific silver fir (Abies amabilis (Dougl.) Forbes) canopy. Specifically, we investigated the role of morphological shade acclimation in the determination of resource use efficiency, which is claimed to be optimal when the distribution of nitrogen within the canopy is directly proportional to the distribution of intercepted photosynthetically active radiation (PAR). Shoots were collected from different heights in the crowns of trees representing four different size classes. A new method was developed to estimate seasonal light interceptance (SLI, intercepted PAR per unit needle area) of the shoots using a model for the directional distribution of above-canopy PAR, measurements of shoot silhouette area and canopy gap fraction in different directions. The ratio SLI/SLI(o), where the reference value SLI(o) represents the seasonal light interceptance of a spherical surface at the shoot location, was used to quantify the efficiency of light capture by a shoot. The ratio SLI/SLI(o) doubled from the top to the bottom of the canopy, mainly as a result of smaller internal shading in shade shoots than in sun shoots. Increased light-capturing efficiency of shade shoots implies that the difference in intercepted light by sun shoots versus shade shoots is much less than the decrease in available light from the upper to the lower canopy. For example, SLI of the five most sunlit shoots was only about 20 times greater than the SLI of the five most shaded shoots, whereas SLI(o) was 40 times greater for sun shoots than for shade shoots. Nitrogen content per unit needle area was about three times higher in sun needles than in shade needles. This variation, however, was not enough to produce proportionality between the amounts of nitrogen and intercepted PAR throughout the canopy.

Journal Article↗

On estimating canopy photosynthesis and stomatal conductance in a deciduous forest with clumped foliage.

The foliage in a fully-leafed deciduous forest canopy is clumped. Consequently, theory indicates that the probability of beam penetration will be estimated more accurately with a model based on the negative binomial distribution than with a model based on the Poisson distribution, incorporating an assumption of a spherical leaf inclination angle distribution. Flux densities of photosynthetically active radiation (PAR) were measured in and above a deciduous forest canopy and were computed with the canopy radiative transfer models based on the negative binomial and Poisson distributions. These radiation values were used to compute canopy photosynthesis and stomatal conductance. Canopy photosynthesis and stomatal conductance, based on the negative binomial model, overestimated values computed from measured PAR profiles by 8 and 9%. respectively. The canopy photosynthesis and stomatal conductance values computed with the spherical Poisson model under-estimated measured values by 17 and 10%, respectively. Thus, the negative binomial radiative transfer model improves estimates of canopy photosynthesis and, to a lesser extent, stomatal conductance, inside a deciduous forest.

Journal Article↗