Effects of kaolin application on light absorption and distribution, radiation use efficiency and photosynthesis of almond and walnut canopies

BACKGROUND AND AIMS: Kaolin applied as a suspension to plant canopies forms a film on leaves that increases reflection and reduces absorption of light. Photosynthesis of individual leaves is decreased while the photosynthesis of the whole canopy remains unaffected or even increases. This may result from a better distribution of light within the canopy following kaolin application, but this explanation has not been tested. The objective of this work was to study the effects of kaolin application on light distribution and absorption within tree canopies and, ultimately, on canopy photosynthesis and radiation use efficiency. METHODS: Photosynthetically active radiation (PAR) incident on individual leaves within the canopy of almond (Prunus dulcis) and walnut (Juglans regia) trees was measured before and after kaolin application in order to study PAR distribution within the canopy. The PAR incident on, and reflected and transmitted by, the canopy was measured on the same day for kaolin-sprayed and control trees in order to calculate canopy PAR absorption. These data were then used to model canopy photosynthesis and radiation use efficiency by a simple method proposed in previous work, based on the photosynthetic response to incident PAR of a top-canopy leaf. KEY RESULTS: Kaolin increased incident PAR on surfaces of inner-canopy leaves, although there was an estimated 20 % loss in PAR reaching the photosynthetic apparatus, due to increased reflection. Assuming a 20 % loss of PAR, modelled photosynthesis and photosynthetic radiation use efficiency (PRUE) of kaolin-coated leaves decreased by only 6.3 %. This was due to (1) more beneficial PAR distribution within the kaolin-sprayed canopy, and (2) with decreasing PAR, leaf photosynthesis decreases less than proportionally, due to the curvature of the photosynthesis response-curve to PAR. The relatively small loss in canopy PRUE (per unit of incident PAR), coupled with the increased incident PAR on the leaf surface on inner-canopy leaves, resulted in an estimated increase in modelled photosynthesis of the canopy (+9 % in both walnut and almond). The small loss in PRUE (per unit of incident PAR) resulted in an increase in radiation use efficiency per unit of absorbed PAR, which more than compensated for the minor (7 %) reduction in canopy PAR absorption. CONCLUSIONS: The results explain the apparently contradictory findings in the literature of positive or no effects of kaolin applications on canopy photosynthesis and yield, despite the decrease in photosynthesis by individual leaves when measured at the same PAR.     

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.     

Estimating photosynthetic radiation use efficiency using incident light and photosynthesis of individual leaves

It has been theorized that photosynthetic radiation use efficiency (PhRUE) over the course of a day is constant for leaves throughout a canopy if leaf nitrogen content and photosynthetic properties are adapted to local light so that canopy photosynthesis over a day is optimized. To test this hypothesis, 'daily' photosynthesis of individual leaves of Solanum melongena plants was calculated from instantaneous rates of photosynthesis integrated over the daylight hours. Instantaneous photosynthesis was estimated from the photosynthetic responses to photosynthetically active radiation (PAR) and from the incident PAR measured on individual leaves during clear and overcast days. Plants were grown with either abundant or scarce N fertilization. Both net and gross daily photosynthesis of leaves were linearly related to daily incident PAR exposure of individual leaves, which implies constant PhRUE over a day throughout the canopy. The slope of these relationships (i.e. PhRUE) increased with N fertilization. When the relationship was calculated for hourly instead of daily periods, the regressions were curvilinear, implying that PhRUE changed with time of the day and incident radiation. Thus, linearity (i.e. constant PhRUE) was achieved only when data were integrated over the entire day. Using average PAR in place of instantaneous incident PAR increased the slope of the relationship between daily photosynthesis and incident PAR of individual leaves, and the regression became curvilinear. The slope of the relationship between daily gross photosynthesis and incident PAR of individual leaves increased for an overcast compared with a clear day, but the slope remained constant for net photosynthesis. This suggests that net PhRUE of all leaves (and thus of the whole canopy) may be constant when integrated over a day, not only when the incident PAR changes with depth in the canopy, but also when it varies on the same leaf owing to changes in daily incident PAR above the canopy. The slope of the relationship between daily net photosynthesis and incident PAR was also estimated from the photosynthetic light response curve of a leaf at the top of the canopy and from the incident PAR above the canopy, in place of that measured on individual leaves. The slope (i.e. net PhRUE) calculated in this simple way did not differ statistically from that calculated using data from individual leaves.     

A simple method to estimate photosynthetic radiation use efficiency of canopies

BACKGROUND AND AIMS: Photosynthetic radiation use efficiency (PhRUE) over the course of a day has been shown to be constant for leaves throughout a general canopy where nitrogen content (and thus photosynthetic properties) of leaves is distributed in relation to the light gradient. It has been suggested that this daily PhRUE can be calculated simply from the photosynthetic properties of a leaf at the top of the canopy and from the PAR incident on the canopy, which can be obtained from weather-station data. The objective of this study was to investigate whether this simple method allows estimation of PhRUE of different crops and with different daily incident PAR, and also during the growing season. METHODS: The PhRUE calculated with this simple method was compared with that calculated with a more detailed model, for different days in May, June and July in California, on almond (Prunus dulcis) and walnut (Juglans regia) trees. Daily net photosynthesis of 50 individual leaves was calculated as the daylight integral of the instantaneous photosynthesis. The latter was estimated for each leaf from its photosynthetic response to PAR and from the PAR incident on the leaf during the day. KEY RESULTS: Daily photosynthesis of individual leaves of both species was linearly related to the daily PAR incident on the leaves (which implies constant PhRUE throughout the canopy), but the slope (i.e. the PhRUE) differed between the species, over the growing season due to changes in photosynthetic properties of the leaves, and with differences in daily incident PAR. When PhRUE was estimated from the photosynthetic light response curve of a leaf at the top of the canopy and from the incident radiation above the canopy, obtained from weather-station data, the values were within 5 % of those calculated with the more detailed model, except in five out of 34 cases. CONCLUSIONS: The simple method of estimating PhRUE is valuable as it simplifies calculation of canopy photosynthesis to a multiplication between the PAR intercepted by the canopy, which can be obtained with remote sensing, and the PhRUE calculated from incident PAR, obtained from standard weather-station data, and from the photosynthetic properties of leaves at the top of the canopy. The latter properties are the sole crop parameters needed. While being simple, this method describes the differences in PhRUE related to crop, season, nutrient status and daily incident PAR.     

Spectral properties of plant leaves pertaining to urban landscape design of broad-spectrum solar ultraviolet radiation reduction

Human exposure to harmful ultraviolet (UV) radiation has important public health implications. Actual human exposure to solar UV radiation depends on ambient UV irradiance, and the latter is influenced by ground reflection. In urban areas with higher reflectivity, UV exposure occurs routinely. To discover the solar UV radiation regulation mechanism of vegetation, the spectral reflectance and transmittance of plant leaves were measured with a spectrophotometer. Typically, higher plants have low leaf reflectance (around 5%) and essentially zero transmittance throughout the UV region regardless of plant species and seasonal change. Accordingly, incident UV radiation decreases to 5% by being reflected and is reduced to zero by passing through a leaf. Therefore, stratified structures of vegetation are working as another terminator of UV rays, protecting whole terrestrial ecosystems, while vegetation at waterfronts contributes to protect aquatic ecosystems. It is possible to protect the human population from harmful UV radiation by urban landscape design of tree shade and the botanical environment. Even thin but uniformly distributed canopy is effective in attenuating UV radiation. To intercept diffuse radiation, UV screening by vertical structures such as hedges should be considered. Reflectivity of vegetation is around 2%, as foliage surfaces reduce incident UV radiation via reflection, while also eliminating it by transmittance. Accordingly, vegetation reduces incident UV radiation to around 2% by reflection. Vegetation influence on ambient UV radiation is broad-spectrum throughout the UV region. Only trees provide cool UV protective shade. Urban landscapes aimed at abating urban heat islands integrated with a reduction of human UV over-exposure would contribute to mitigation of climate change.     

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.     

Modeling the vertical foliage distribution of an individual <Emphasis Type="Italic">Castanopsis cuspidata</Emphasis> (Thunb.) Schottky,a dominant broad-leaved tree in Japanese warm-temperate forest

The vertical foliage distribution of Castanopsis cuspidata (Thunb.) Schottky was examined in trees of various sizes to clarify its variation in relation to tree size and the light environment in a stand. As indices of these parameters, we analyzed crown social position (CSP: percent of stand height) and specific leaf area (SLA). The vertical foliage distribution of trees was expressed by a Weibull function. The variation in the vertical foliage distribution of C. cuspidata could be categorized into three types using crown social position and light environment. In the first type, leaves were concentrated to the top 20% of the tree; such trees are canopy trees that can receive full sunlight. The second type had a large relative crown depth and an asymmetric distribution with the maximum foliage located near the top of the tree; such trees are suppressed trees whose crowns do not receive sufficient light. The third type had a large relative crown depth and a symmetric distribution; such trees occur in high light environments, although their crowns are in the understory layer. The differences in the vertical foliage distribution are related to the strategies used to capture light. Multiple regression analysis showed that CSP and SLA at the top layer of the tree explained successive changes in the vertical foliage distribution. These results will contribute to scaling-up the vertical foliage distribution to the community level in pure stands of C. cuspidata using an individual-based model.     

Angle distribution of foliage in individual Chamaecyparis obtusa canopies and effect of angle on diffuse light penetration

 The vertical distribution of foliage angle and area of three Chamaecyparis obtusa trees was determined by the triangle method, which calculates foliage geometry using measured coordinates of the leaf ”corners”, in a 43-year-old plantation in central Japan. Vertical distribution patterns of leaf area were different depending on tree size, but the boundary heights, which divide the canopy into sunlit and shaded parts, were similar in the three sample trees. The value of the average foliage angle [I(Z)] at a given depth (Z) from the tip of the stem decreased continually from the upper to lower layers within the canopy. The vertical patterns of changes in I(Z) were different among the three trees, but could be expressed by the following allometric equation as a function of depth.

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

Colonization Strategies of Two Liana Species in a Tropical Dry Forest Canopy

Lianas impose intense resource competition for light in the upper forest canopy by displaying dense foliage on top of tree crowns. Using repeated access with a construction crane, we studied the patterns of canopy colonization of the lianas Combretum fruticosum and Bonamia trichantha in a Neotropical dry forest in Panama. Combretum fruticosum flushed leaves just before the rainy season, and its standing leaf area quickly reached a peak in the early rainy season (May–June). In contrast, B. trichantha built up foliage area continuously throughout the rainy season and reached a peak in the late rainy season (November). Both species displayed the majority of leaves in full sun on the canopy surface, but C. fruticosum displayed a greater proportion of leaves (26%) in more shaded microsites than B. trichantha (12%). Self-shading within patches of liana leaves within the uppermost 40–50 cm of the canopy reduced light levels measured with photodiodes placed directly on leaves to 4–9 percent of light levels received by sun leaves. Many leaves of C. fruticosum acclimated to shade within a month following the strongly synchronized leaf flushing and persisted in deep shade. In contrast, B. trichantha produced short-lived leaves opportunistically in the sunniest locations. Species differences in degree of shade acclimation were also evident in terms of structural (leaf mass per area, and leaf toughness) and physiological characters (nitrogen content, leaf life span, and light compensation point). Contrasting leaf phenologies reflect differences in light exploitation and canopy colonization strategies of these two liana species.     

The alpha-tocopherol content of leaves of pedunculate oak (Quercus robur L.)--variation over the growing season and along the vertical light gradient in the canopy

This study was performed in order to investigate whether the actual requirement for defence against photo-oxidative stress is reflected by the alpha-tocopherol (alpha-Toco) content in leaves of pedunculate oak (Quercus robur L.). Antioxidants and pigments were quantified in leaves that were collected on six days between May and September 2000 in a mixed pine/oak forest at canopy positions differing in light environment. Pools of hydrophilic antioxidants and photo-protective xanthophyll cycle pigments (V + A + Z) reflected the anti-oxidative demand, as these pools increased with the average light intensity to which the leaves were acclimated. The photo-protective demand was not the determinant of the alpha-Toco content of oak leaves, as (1) foliage of a young oak, exposed to low light levels in the understorey, contained higher amounts of this lipophilic antioxidant than leaves sampled from semimature oaks at canopy positions with a similar light environment, and (2) a strong increase in the alpha-Toco content over the growing season was detected at each investigated crown position, whereas the V + A + Z pool did not show a concomitant accumulation during leaf ageing. The rate of alpha-Toco accumulation differed distinctly between samples taken at different canopy positions.     

The effects of the spatial pattern of defoliation on regrowth of a tussock grass

Summary The spatial pattern of foliage removal from a tussock grass can influence regrowth through effects on daily carbon gain (CER_d). This field study examined the extent to which tussock photosynthetic responses to different defoliation patterns were associated with changes in whole-canopy attributes (e.g., foliage age structure, canopy light microclimate). During the spring growing season, 60% of the green foliage area was removed from individual Agropyron desertorum tussocks with scissors in different spatial patterns. These patterns represented extremes of defoliation patterns that might be inflicted by natural herbivores. Tussock photosynthesis (per unit foliage area) at high light (2000 mol photons m^–2 s^–1 between 400 and 700 nm; P₂₀₀₀) increased following clipping with all defoliation patterns. The increases in P₂₀₀₀ were greater when leaves were removed from low in the tussock (older leaves) than if leaves high in the canopy (younger leaves) were removed. These relative changes of P₂₀₀₀ among clipping patterns paralleled the responses of CER_d and regrowth from an earlier study. Furthermore, the changes in P₂₀₀₀ corresponded with increases in the proportion of foliage within the tussocks that was directly illuminated at midday. The greater photosynthesis of tussocks after lower-leaf removal was directly related to a higher proportion of younger foliage and a smaller fraction of foliage shaded within the tussock. In a dense canopy, such as these grass tussocks, the influence of defoliation on whole-canopy attributes may be of primary importance to whole-plant photosynthetic responses.     

Coping with branch excision when measuring leaf net photosynthetic rates in a lowland tropical forest

Measuring leaf gas exchange from canopy leaves is fundamental for our understanding of photosynthesis and for a realistic representation of carbon uptake in vegetation models. Since canopy leaves are often difficult to reach, especially in tropical forests with emergent trees up to 60 m at remote places, canopy access techniques such as canopy cranes or towers have facilitated photosynthetic measurements. These structures are expensive and therefore not very common. As an alternative, branches are often cut to enable leaf gas exchange measurements. The effect of branch excision on leaf gas exchange rates should be minimized and quantified to evaluate possible bias. We compared light-saturated leaf net photosynthetic rates measured on excised and intact branches. We selected branches positioned at three canopy positions, estimated relative to the top of the canopy: upper sunlit foliage, middle canopy foliage, and lower canopy foliage. We studied the variation of the effects of branch excision and transport among branches at these different heights in the canopy. After excision and transport, light-saturated leaf net photosynthetic rates were close to zero for most leaves due to stomatal closure. However, when the branch had acclimated to its new environmental conditions—which took on average 20 min—light-saturated leaf net photosynthetic rates did not significantly differ between the excised and intact branches. We therefore conclude that branch excision does not affect the measurement of light-saturated leaf net photosynthesis, provided that the branch is recut under water and is allowed sufficient time to acclimate to its new environmental conditions.     

Ecology And Growth Habit Of Laveineopteris: A Gymnosperm from the Late Carboniferous Tropical Rain Forests

Cyclopterid leaves were borne on the same Late Carboniferous medullosalean plants that bore the pinnate fronds currently known as Laveineopteris . They were morphologically and anatomically different from the pinnate foliage, and presumably were also physiologically different. The cyclopterids are here interpreted as having been shade leaves and the pinnate foliage sun leaves. The juvenile Laveineopteris plant probably consisted of a monopole sapling bearing only cyclopterid leaves, which optimized growth in canopy shade conditions. On reaching the canopy level, the plant produced a crown of pinnate sun leaves. Cyclopterids were also borne epiphyllously in the proximal parts of the pinnate fronds; this may have represented a transitional light–level between canopy and subcanopy conditions. Thus, in this reconstruction, the cyclopterid leaves were complementary to the pinnate foliage of the adult Laveineopteris plant, and played a key role in its growth habit and ecology.     

Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

Geometrical similarity analysis of photosynthetic light response curves, light saturation and light use efficiency

Light absorption and use efficiency (LAUE mol mol⁻¹, daily gross photosynthesis per daily incident light) of each leaf depends on several factors, including the degree of light saturation. It is often discussed that upper canopy leaves exposed to direct sunlight are fully light-saturated. However, we found that upper leaves of three temperate species, a heliophytic perennial herb Helianthus tuberosus, a pioneer tree Alnus japonica, and a late-successional tree Fagus crenata, were not fully light-saturated even under full sunlight. Geometrical analysis of the photosynthetic light response curves revealed that all the curves of the leaves from different canopy positions, as well as from the different species, can be considered as different parts of a single non-rectangular hyperbola. The analysis consistently explained how those leaves were not fully light-saturated. Light use optimization models, called big leaf models, predicted that the degree of light saturation and LAUE are both independent of light environment. From these, we hypothesized that the upper leaves should not be fully light-saturated even under direct sunlight, but instead should share the light limitation with the shaded lower-canopy leaves, so as to utilize strong sunlight efficiently. Supporting this prediction, within a canopy of H. tuberosus, both the degree of light saturation and LAUE were independent of light environment within a canopy, resulting in proportionality between the daily photosynthesis and the daily incident light among the leaves.     

Effect of light interception by non-photosynthetic organs on canopy photosynthetic production

A canopy photosynthesis model was derived on the assumption that the light diminution within a canopy is caused by both leaves and non-photosynthetic organs. The light diminution by leaves and that by non-photosynthetic organs were taken into account separately in the Lambert-Beer equation of light extinction. The light flux density on the leaf surface at each depth was evaluated from the leaf's share of light. The light flux density on the leaf surface thus obtained was incorporated into the Monsi-Saeki model of canopy photosynthesis. The proposed model was applied for estimating gross canopy photosynthesis in a 19-year-oldLarix leptolepis plantation where 38% of the light diminution was due to non-photosynthetic organs. The daily canopy photosynthesis on one summer day calculated using the present model was about 22% less than that calculated by the conventional Monsi-Saeki model, in which light interception by non-photosynthetic organs is neglected. The degree of such reduction in canopy photosynthesis through shading by non-photosynthetic organs was assessed in relation to parameters affecting light extinction, leaf photosynthetic characteristics, and light regime above the canopy.     

Variation in crown light utilization characteristics among tropical canopy trees

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

The effects of light acclimation during and after foliage expansion on photosynthesis ofAbies amabilis foliage within the canopy

Variation in the photosynthetic function ofAbies amabilis foliage within a canopy was examined and related to three different processes that affect foliage function: foliage aging, sun-shade acclimation that occurred while foliage was expanding, and reacclimation after expansion was complete. Foliage produced in the sun had higher photosynthesis at light saturation (A _max, mol·m^-2·s^-1), dark respiration (mol·m^-2·s^-1), nitrogen content (g·m^-2), chlorophyll content (g·m^-2), and chlorophylla:b ratio, and a lower chlorophyll to nitrogen ratio (chl:N), than foliage produced in the shade. As sun foliage becomes shaded, it becomes physiologically similar to shade foliage, even though it still retains a sun morphology. Shaded sun foliage exhibited lowerA _max, dark respiration, nitrogen content, and chlorophylla:b ratio, and a higher chl:N ratio than sun foliage of the same age remaining in the open. However, shaded sun foliage had a higher chlorophyll content than sun foliage remaining in the open, even though true shade foliage had a lower chlorophyll content than sun foliage. This anomaly arises because as sun foliage becomes shaded, it retains a higher nitrogen content than shade foliage in a similar light environment, but the two forms have similar chl:N ratios. Within the canopy, most physiological indicators were more strongly correlated with the current light environment than with foliage age or leaf thickness, with the exception of chlorophyll content.A _max decreased significantly with both decreasing current light environment of the foliage and increasing foliage age. The same trend with current light and age was found for the chlorophylla:b ratio. Foliage nitrogen content also decreased with a decrease in current light environment, but no distinct pattern was found with foliage age. Leaf thickness was also important for predicting leaf nitrogen content: thicker leaves had more nitrogen than thinner leaves regardless of light environment or age. The chl:N ratio had a strong negative correlation with the current light environment, and, as with nitrogen content, no distinct pattern was found with foliage age. Chlorophyll content of the foliage was not well correlated with any of the three predictor variables: current light environment, foliage age or leaf thickness. On the other hand, chlorophyll content was positively correlated with the amount of nitrogen in a leaf, and once nitrogen was considered, the current light environment was also highly significant in explaining the variation in chlorophyll content. It has been suggested that the redistribution of nitrogen both within and between leaves is a mechanism for photosynthetic acclimation to the current light environment. Within theseA. amabilis canopies, both leaf nitrogen and the chl:N ratio were strongly correlated with the current light environment, but only weakly with leaf age, supporting the idea that changing light is the driving force for the redistribution of nitrogen both within and between leaves. Thus, our results support previous theories on nitrogen distribution and partitioning. However,A _max was significantly affected by both foliage age and the current light environment, indicating that changes in light alone are not enough to explain changes inA _max with time.     

不同光环境下桂花幼苗的构型差异

通过对不同光照条件下桂花幼苗的冠形、分枝率、叶片在树冠中的空间分布等特征进行研究,结果表明桂花幼苗构型发生了明显的可塑性适应:其树冠对光照条件的变化有显著的可塑性响应。在林隙中的幼苗受光的间歇性影响,总体分枝率明显小于全光、林冠下的幼苗分枝率。全光的幼苗叶片集中于二级枝,叶片长度和叶片面积相对较小,对光照利用充分;而林隙中的幼苗叶片集中于一级枝,避免处于植冠内侧受到遮蔽,表现出较大的叶片长度和叶面积;林冠下的叶片较均匀分布在一、二级枝上,叶片总数量较少,枝条高生长较全光下明显。幼苗在总体分枝格局中表现出独自的特点,即强光环境下产生短枝和高分枝率,在适度庇荫条件下产生长枝及低分枝率,在强度庇荫条件下以较长枝和较高分枝率来同时满足高生长和横向生长的需求。