Photosynthetic responses to light variation in rainforest species

Summary The dependence of net carbon gain during lightflecks (artificial sunflecks) on leaf induction state, lightfleck duration, lightfleck photosynthetic photon flux density (PFD), and the previous light environment were investigated in A. macrorrhiza and T. australis, two Australian rainforest species. The photosynthetic efficiency during lightflecks was also investigated by comparing observed values of carbon gain with predicted values based on steady-state CO₂ assimilation rates. In both species, carbon gain and photosynthetic efficiency increased during a series of five 30-or 60-s lightflecks that followed a long period of low light; efficiency was linearly related to leaf induction state.In fully-induced leaves of both species, efficiency decreased and carbon gain increased with lightfleck duration. Low-light grown A. macrorrhiza had greater efficiency than predicted based on steady-state rates (above 100%) for lightflecks less than 40 s long, whereas leaves grown in high light had efficiencies exceeding 100% only during 5-s lightflecks. The efficiency of leaves of T. australis ranged from 58% for 40-s lightflecks to 96% for 5-s lightflecks.In low-light grown leaves of A. macrorrhiza, photosynthetic responses to lightflecks below 120 mol m^-2 s^-1 were not affected significantly by the previous light level. However, during lightflecks at 530 mol m^-2 s^-1, net carbon gain and photosynthetic efficiency of leaves previously exposed to low light levels were significantly reduced relative to those of leaves previously exposed to 120 and 530 mol m^-2 s^-1.These results indicate that, in shade-tolerant species, net carbon gain during sunflecks can be enhanced over values predicted from steady-state CO₂ assimilation rates. The degree of enhancement, if any, will depend on sunfleck duration, previous light environment, and sunfleck PFD. In forest understory environments, the temporal pattern of light distribution may have far greater consequences for leaf carbon gain than the total integrated PFD.Supported by National Science Foundation Grant BSR 8217071 and USDA Grant 85-CRCR-1-1620     

A preliminary study on the photosynthetic induction response ofQuercus serrata seedlings

To understand the sunfleck utilization of leaves in heliophilic trees within grass canopies, we studied the photosynthetic induction response ofQuercus serrata seedlings grown for 5 months in different microsites in aMiscanthus sinensis canopy. Two phases, a rapid increase in CO₂ uptake and a following slow increase, were recognizabie in the time course of CO₂ uptake in response to an increase of photon flux densities (PFD). When the preceding period of low light became shorter, the period of the two phases became shorter. The capacity of response to a sudden light increase was evaluated by the relative photosynthetic induction efficiency (RPIE) defined as the ratio of integrated carbon gain measured to that calculated by assuming that a steady-state assimilation would be achieved instantaneously after the light increase. RPIEs estimated were negatively related to potential sunfleck PFD of microsite. The leaves of the seedlings grown in the microsite with a lower sunfleck PFD and a shorter sunfleck duration showed a more rapid response to a sudden increase of light. These findings suggest that the leaves ofQ. serrata seedlings growing under a lower sunfleck PFD are able to increase the photosynthetic capacity more rapidly to an increase of PFD.     

Steady-state and dynamic photosynthetic response of Adenocaulon bicolor (Asteraceae) in its redwood forest habitat

Summary The gas exchange characteristics under steadystate and transient light conditions were determined for a redwood forest understory herb Adenocaulon bicolor, that depends on use of sunflecks for a large fraction of its daily carbon gain. Measurements under steady-state conditions indicated that this species has photosynthetic characteristics that are typical for understory plants. The mean light-saturated assimilation rate was 5.26 mol CO₂ m^-2 s^-1; the light saturation and compensation occurred at 243 and 2 mol photons m^-2 s^-1, respectively. This light compensation point was much less than the photon flux density under diffuse light in the understory so that positive assimilation could be maintained throughout the day. When leaves that had been in diffuse light for at least 2 h were exposed to a sudden increase in PFD to saturating levels, 10–30 min were required for both assimilation and stomatal conductance to reach maximum values. Calculation of intercellular CO₂ pressures, however, suggest that for the first 10 min after the light increase, biochemical factors were responsible for most of the increase in assimilation. Thereafter stomatal opening caused a further increase in assimilation that was no more than 25% of the total. When fully induced leaves were returned to low light, induction was rapidly lost even though stomatal conductance decreased only slowly. This rapid loss of induction limited the capacity of A. bicolor to use sunflecks after low light periods that lasted longer than 1–2 min. However, during periods when sunflecks are more frequent there is probably little loss of induction. Under these conditions, sunflecks are used with high efficiency for assimilation.     

Heterogeneity of Light Availability and its Effects on Simulated Carbon Gain of Tree Leaves in a Small Gap and the Understory in a Tropical Rain Forest1

To clarify the small-scale heterogeneity of light regimes in a rain forest, photosynthetic photon flux density (PFD) was measured at 1-min intervals during six days at 12 microsites in each of two plots, a small gap and an understory in Pasoh Forest Reserve, Peninsular Malaysia. Frequency distribution of microsite PFD was unimodal with the peak value between 16 and 32 μmol/m²/sec in the small gap, but between 8 and 16 μmol/m²/sec in the understory. In the small gap, PFD was more variable among microsites; total daily PFD and daily sunfleck PFD exceeding 10 μmol/ m²/sec tended to be higher (P <0.05; t-test) compared to those in the understory. Sunfleck PFD exceeding 50 μmol/ m²/sec, however, showed no difference between the two plots. Diffuse PFD transmittance, defined as the ratio of PFD in the forest to that measured at 43 m above ground during the periods 0800-0810 and 1750-1800 h, was significantly higher in the small gap than in the understory plot. Diffuse PFD transmittance was also positively correlated with microsite total daily PFD. To examine the effects of the subtle heterogeneity of light regimes on leaf carbon gain, we simulated carbon gain by sun and shade leaves in a typical shade-tolerant species, Brosimum aticastrum Sw. (Moraceae). Despite the similarity in total daily PFD, total daily carbon gain was considerably higher in the gap than in the understory for both sun and shade leaves. This study suggests that frequency distribution of PFD is critical in describing microsite PFD regimes and determining leaf carbon gain in the tropical forest floor.     

Relative enhancement of photosynthesis and growth at elevated CO2 is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survivorship of dipterocarp seedlings in the deeply shaded understorey of South‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. To investigate the effect of elevated CO₂ upon photosynthesis and growth under sunflecks, seedlings of Shorealeprosula were grown in controlled environment conditions at ambient or elevated CO₂. Equal total daily photon flux density (PFD) (～7·7 mol m^?2 d^?1) was supplied as either uniform irradiance (～170 µmol m^?2 s^?1) or shade/fleck sequences (～30 µmol m^?2 s^?1/～525 µmol m^?2 s^?1). Photosynthesis and growth were enhanced by elevated CO₂ treatments but lower under flecked irradiance treatments. Acclimation of photosynthetic capacity occurred in response to elevated CO₂ but not flecked irradiance. Importantly, the relative enhancement effects of elevated CO₂ were greater under sunflecks (growth 60%, carbon gain 89%) compared with uniform irradiance (growth 25%, carbon gain 59%). This was driven by two factors: (1) greater efficiency of dynamic photosynthesis (photosynthetic induction gain and loss, post‐irradiance gas exchange); and (2) photosynthetic enhancement being greatest at very low PFD. This allowed improved carbon gain during both clusters of lightflecks (73%) and intervening periods of deep shade (99%). The relatively greater enhancement of growth and photosynthesis at elevated CO₂ under sunflecks has important potential consequences for seedling regeneration processes and hence forest structure and composition.     

Fluctuation of photosynthetic photon flux density within aMiscanthus sinensis canopy

Detailed measurements of diurnal variations in photosynthetic photon flux density (PPFD) were made at seven locations within the canopy of aMiscanthus sinensis grassland to evaluate the light conditions of microsites for heliophilic tree seedlings. Multiple regression analysis revealed that the short-term light fluctuation on a clear day was highly dependent on the wind speed and solar elevation angle, whereas on a cloudy day it was mainly determined by the PPFD incident from above the canopy. The relative PPFD at 40 cm aboveground varied from 0.065 to 0.252, depending on sky conditions and the sensor's position in relation to clumped patches ofM. sinensis. On a clear day, the proportion of PPFD readings above 100 μmol·m⁻²·s⁻¹ contributed by sunflecks ranged between 25.4% and 82.0%. Computer simulation showed that the contribution of sunflecks to the daily carbon gain ofQuercus serrata seedlings may range from 11% to 65%. The mean relative PPFD measured under diffuse light conditions was linearly related to the daily total PPFD and the daily carbon gain by single leaves ofQ. serrata seedlings. This suggests that the relative PPFD under diffuse light conditions provides an appropriate measure of site-specific light availability within a grass canopy.     

Adaptive radiation of photosynthetic physiology in the Hawaiian lobeliads: dynamic photosynthetic responses

Hawaiian lobeliads have radiated into habitats from open alpine bogs to densely shaded rainforest interiors, and show corresponding adaptations in steady-state photosynthetic light responses and associated leaf traits. Shaded environments are not uniformly dark, however, but punctuated by sunflecks that carry most of the photosynthetically active light that strikes plants. We asked whether lobeliads have diversified in their dynamic photosynthetic light responses and how dynamic responses influence daily leaf carbon gain. We quantified gas exchange and dynamic light regimes under field conditions for ten species representing each major Hawaiian sublineage. Species in shadier habitats experienced shorter and less numerous sunflecks: average sunfleck length varied from 1.4 ± 1.7 min for Cyanea floribunda in shaded forest understories to 31.2 ± 2.1 min for Trematolobelia kauaiensis on open ridges. As expected, the rate of photosynthetic induction increased significantly toward shadier sites, with assimilation after 60 s rising from ca. 30% of fully induced rates in species from open environments to 60% in those from densely shaded habitats. Uninduced light use efficiency—actual photosynthesis versus that expected under steady-state conditions—increased from 10 to 70% across the same gradient. In silico transplants—modeling daily carbon gain using one species’ photosynthetic light response in its own and other species’ dynamic light regimes—demonstrated the potential adaptive nature of species differences: understory Cyanea pilosa in its light regimes outperformed gap-dwelling Clermontia parviflora, while Clermontia in its light regimes outperformed Cyanea. The apparent crossover in daily photosynthesis occurred at about the same photon flux density where dominance shifts from Cyanea to Clermontia in the field. Our results further support our hypothesis that the lobeliads have diversified physiologically across light environments in Hawaiian ecosystems and that those shifts appear to maximize the carbon gain of each species in its own environment. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal variations of microsite light availability within aMiscanthus sinensis canopy

Photosynthetic photon flux density (PPFD) at 15 cm above the ground was measured at 20 microsites in gaps and grass patches within aMiscanthus sinensis Anderss community at 10 s intervals during 5 days every month from May to September 1989. Microsite light availability, which was characterized by daily total PPFD, sunfleck PPFD (PPFD above a threshold value of 50 or 400 μmol m⁻² s⁻¹) and the diffuse site factor, showed evident seasonal changes, with a marked reduction between June and July due to the rapid growth of the grass canopy. The monthly median value of daily total PPFD among the microsites decreased from 10.3 mol m⁻² day⁻¹ in May to 0.77 mol m⁻² day⁻¹ in September, with a reduction in the diffuse site factor from 31 to 4%. During the summer, the median value of the total time of sunflecks exceeding 50 μmol m⁻² s⁻¹ contributed 7–18% of measurement time, but the contribution of these sunflecks to daily total PPFD ranged from 29 to 59%. There was considerable microsite variation in light availability throughout the measurement period. Rank correlation analysis revealed that some microsites, such as those in gaps, consistently received more total PPFD, more sunfleck PPFD and had a higher diffuse site factor than those in grass patches. The diffuse site factor had a linearly positive relationship with daily total PPFD and total sunfleck PPFD for the 20 microsites during the measurement period, confirming that the diffuse site factor is a useful index for microsite light availability withinM. sinensis canopies.     

Light environment and carbon gain of understory herbs associated with sunflecks in a warm temperate deciduous forest in Japan

Seasonal variation in the light environment on the forest floor of a deciduous forest was investigated with special reference to sunflecks. Diurnal variations and seasonal changes in frequency and irradiation period of the sunflecks (sunfleck duration) were measured. The hourly total sunfleck duration varied seasonally; that is, 30–40 min in spring and autumn and about 15–20 min in summer. There was no large variation in the hourly sunfleck duration during daytime hours (from 9.00 to 15.00 h). The emergence frequency of sunflecks was 1.3–4.8 per h with two peaks, one in the morning and one in the afternoon. The mean duration of a sunfleck, however, showed a characteristic daily pattern with a peak around noon. Sunfleck duration was long around noon, ranging from 12 to 18 min, and short around 10.00 and 14.00 h, ranging from 6 to 10 min. Using the light photosynthesis curves ofPyrola japonica andSyneilesis palmata (Koizumi & Oshima 1985), the contribution of sunflecks to the dry matter production of these understory species was evaluated. It was shown that the sunflecks contributed 7–10% of the carbon gain inS. palmata, but only 2–3% of that inP. japonica.     

Modeling dynamic understory photosynthesis of contrasting species in ambient and elevated carbon dioxide

Dynamic responses of understory plants to sunflecks have been extensively studied, but how much differences in dynamic light responses affect daily photosynthesis (A_day) is still the subject of active research. Recent models of dynamic photosynthesis have provided a quantitative tool that allows the critical assessment of the importance of these sunfleck responses on A_day. Here we used a dynamic photosynthesis model to assess differences in four species that were growing in ambient and elevated CO₂. We hypothesized that Liriodendron tulipifera, a species with rapid photosynthetic induction gain and slow induction loss, would have the least limitations to sunfleck photosynthesis relative to the other three species (Acer rubrum, Cornus florida, Liquidambar styraciflua). As a consequence, L. tulipifera should have the highest A_day in an understory environment, despite being the least shade tolerant of the species tested. We further hypothesized that daily photosynthetic enhancement by elevated CO₂ would differ from enhancement levels observed during light-saturated, steady-state measurements. Both hypotheses were supported by the model results under conditions of low daily photosynthetic photon flux density (PFD; <3% of the above-canopy PFD). However, under moderate PFD (10-20% of the above-canopy PFD), differences in dynamic sunfleck responses had no direct impact on A_day for any of the species, since stomatal and photosynthetic induction limitations to sunfleck photosynthesis were small. Thus, the relative species ranking in A_day under moderate PFD closely matched their rankings in steady-state measurements of light-saturated photosynthesis. Similarly, under elevated CO₂, enhancement of modeled A_day over A_day at ambient CO₂ matched the enhancement measured under light saturation. Thus, the effects of species-specific differences in dynamic sunfleck responses, and differences in elevated CO₂ responses of daily photosynthesis, are most important in marginal light environments.     

Photosynthetic responses of two heliotropic legumes from contrasting habitats

Abstract Photosynthetic responses to light, temperature and leaf-to-air water vapour concentration deficit for Strophostyles helvola (L.) Ell. in an open beach site, and Amphicarpa bracteata (L.) Ell. in two deciduous forest sites were quantified. Photosaturated rates of net CO₂ assimilation were 52.1 ± 4.6, 11.0 ± 1.6 and 4.1 ± 0.3 μmol m^?2s^?2 for plants in beach, roadside and closed canopy sites, respectively. In terms of photosynthesis, plants in the beach site were more tolerant of higher leaf temperatures and water vapour concentration deficits than were plants in forested sites. Heliotropic leaf movements in the beach site reduced calculated total daily transpiration by 2%, increased total daily carbon gain by 8% and reduced the transpiration ratio by 9% relative to an horizontal leaf. During long-term sunflecks in forested sites, heliotropism reduced transpiration by 15%, increased carbon gain by 71% and reduced the transpiration ratio by 50% relative to an horizontal leaf. We hypothesize that heliotropic leaf movements in mesic, high-light, low-nitrogen habitats may increase carbon return on nitrogen investment in photosynthesis, while heliotropism in canopy gaps may represent a morphological mechanism to reduce damage to low-light acclimated photosynthetic systems during long-term sunflecks.     

High-temperature inhibition of photosynthesis is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survival of dipterocarp seedlings in the understorey of south‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. Field measurements demonstrated that Shorea leprosula seedlings in a rain forest understorey received a high proportion of daily photon flux density at temperatures supra‐optimal for photosynthesis (72% at ≥30 °C, 14% at ≥35 °C). To investigate the effect of high temperatures on photosynthesis during sunflecks, gas exchange and chlorophyll fluorescence measurements were made on seedlings grown in controlled environment conditions either, under uniform, saturating irradiance (approximately 539 µmol m^?2 s^?1) or, shade/fleck sequences (approximately 30 µmol m^?2 s^?1/approximately 525 µmol m^?2 s^?1) at two temperatures, 28 or 38 °C. The rate of light‐saturated photosynthesis, under uniform irradiance, was inhibited by 40% at 38 °C compared with 28 °C. However, during the shade/fleck sequence, photosynthesis was inhibited by 59% at 38 °C compared with 28 °C. The greater inhibition of photosynthesis during the shade/fleck sequence, when compared with uniform irradiance, was driven by the lower efficiency of dynamic photosynthesis combined with lower steady‐state rates of photosynthesis. These results suggest that, contrary to current dogma, sunfleck activity may not always result in significant carbon gain. This has important consequences for seedling regeneration processes in tropical forests as well as for leaves in other canopy positions where sunflecks make an important contribution to total photon flux density.     

A functional analysis of the crown architecture of tropical forest<Emphasis Type="Italic"> Psychotria</Emphasis> species: do species vary in light capture efficiency and consequently in carbon gain and growth?

The crown architectures of 11 Psychotria species native to Barro Colorado Island, Panama were reconstructed from field measurements of leaf and branch geometry with the three-dimensional simulation model Y-plant. The objective was to assess the role of species differences in architecture in light capture and carbon gain in their natural understory environment. When species were grouped according to their putative light environment preference, the shade tolerant species were found to have a small but significantly higher efficiency of light capture for both diffuse and direct light as compared to the light demanding species. Within each grouping, however, there were few significant differences in light capture efficiency among species. The lower efficiencies of light demanding species was due to slightly higher self-shading and slightly lower angular efficiencies. Simulations of whole plant assimilation showed that light demanding species had greater daily assimilation in both direct and diffuse light due to the significantly greater light availability in the sites where light demanding species were found, as compared to those where shade tolerant species occurred. Among light demanding species, the above ground relative growth rate measured over a 1-year period by applying allometric equations for mass versus linear dimensions, was positively correlated with diffuse PFD and with mean daily assimilation estimated from Y-plant. For the shade tolerant plants, there was no significant correlation between RGR and mean daily assimilation or with any measure of light availability, probably because they occurred over a much narrower range of light environments. Overall, the results reveal a strong convergence in light capture efficiencies among the Psychotria species at lower values than previously observed in understory plants using similar approaches. Constraints imposed by other crown functions such as hydraulics and biomechanical support may place upper limits on light capture efficiency.Abbreviations E_a Efficiency of light absorption (dimensionless) - E_adir Efficiency of direct light absorption (dimensionless) - E_adif Efficiency of diffuse light absorption (dimensionless) - D_E Display efficiency (dimensionless) - P_E Projection efficiency (dimensionless) - CosI Mean cosine of incidence (dimensionless) - aLAR_e Effective leaf area ratio (m² g^–1) - A_tot Daily assimilation (mmol m^–2 day^–1) - A_dir Daily assimilation in direct PFD (mmol m^–2 day^–1) - A_dif Daily assimilation in diffuse PFD (mmol m^–2 day^–1)     

Microsite variation in light availability and seedling growth ofQuercus serrata in a temperate pine forest

Spatial heterogeneity in light availability for tree seedlings under the canopy of a temperate pine forest was studied. Six-day measurements at 10-s intervals revealed a great variety in the temporal patterns of photosynthetic photon flux density (PPFD) and histograms among observation days and microsites; mean daily total PPFD relative to full sun varied from 1.5% to 10.4% depending on the microsites. The occurrence and duration of PPFD above 80 μmol m⁻² s⁻¹, which might reflect sunfleck activity, varied greatly among the microsites. However, several simple empirical relationships were found between some parameters characterizing microsite light availability and sunfleck activity; the diffuse site factor was correlated well with other parameters, including daily total PPFD, daily totals and daily summed durations of high PPFD above any examined threshold level, and its contribution to daily total PPFD. Diffuse site factors which were obtained for 700 microsites within an area of 28 m² on three different occasions during the growing season showed high correlations within the microsite. Based on the regressed relationship between the relative growth rate of current-year seedlings ofQuercus serrata and the microsite diffuse site factor and the results of area-survey measurement of the diffuse site factor, an estimation was made of the abundance of potential ‘safe-sites’ for seedling growth of the species; the ‘safe sites’ were estimated to cover 40% and 0% of the total area of the sunny and shady sites of the forest, respectively.     

Analysis of the ��shelter tree-effect�� of natural and exotic forest canopies on the growth of young Podocarpus falcatus trees in southern Ethiopia

Plantations of exotic trees on areas where tropical forest has been clear-felled have been reported to facilitate regrowth of indigenous tree species. This so-called nurse- or shelter tree effect was investigated in a montane semihumid site in southern Ethiopia where plantations of Pinus patula and Eucalyptus saligna grow in close vicinity to the natural Podocarpus falcatus mixed forest. Physiological performance of P. falcatus saplings growing in the exotic and the natural forests was investigated over the year. Compared with the natural forest, photosynthetic carbon gain and growth of the saplings were significantly enhanced under Pinus patula while likewise retarded under Eucalyptus saligna. Diverging effects of the differently dense shelter canopies on the saplings could be traced to differences in the sub-canopy microclimates and particularly to the intensities and temporal distribution of photosynthetic active radiation (PAR). Moisture also played an important role for photosynthetic carbon gain: while the morning patterns of CO₂ uptake were significantly correlated with the patterns of sunflecks, stomatal conductance was the determinant factor in the afternoon. Analysis of the photosynthetic efficiency of sunflecks revealed a lower quantum yield than the basic diffuse PAR intensity. Compared with a theoretically even distribution of the daily PAR, CO₂ uptake under the real light climate was 70% under Pinus and in the natural forest, and 59% under Eucalyptus. Relating growth rates of Podocarpus saplings to photosynthesis the microclimate under Pinus was 2.5 times as effective as that in the natural forest and five times more effective than under Eucalyptus.     

The functional ecology of shoot architecture in sun and shade plants of Heteromeles arbutifolia M. Roem., a Californian chaparral shrub

The functional roles of the contrasting morphologies of sun and shade shoots of the evergreen shrub Heteromeles arbutifolia were investigated in chaparral and understory habitats by applying a three-dimensional plant architecture simulation model, YPLANT. The simulations were shown to accurately predict the measured frequency distribution of photosynthetic photon flux density (PFD) on both the leaves and a horizontal surface in the open, and gave reasonably good agreement for the more complex light environment in the shade. The sun shoot architecture was orthotropic and characterized by steeply inclined (mean = 71^o) leaves in a spiral phyllotaxy with short internodes. This architecture resulted in relatively low light absorption efficiencies (E _A) for both diffuse and direct PFD, especially during the summer when solar elevation angles were high. Shade shoots were more plagiotropic with longer internodes and a pseudo-distichous phyllotaxis caused by bending of the petioles that positioned the leaves in a nearly horizontal plane (mean = 5^o). This shade-shoot architecture resulted in higher E _A values for both direct and diffuse PFD as compared to those of the sun shoots. Differences in E _A between sun and shade shoots and between summer and winter were related to differences in projection efficiencies as determined by leaf and solar angles, and by differences in self shading resulting from leaf overlap. The leaves exhibited photosynthetic acclimation to the sun and the shade, with the sun leaves having higher photosynthetic capacities per unit area, higher leaf mass per unit area and lower respiration rates per unit area than shade leaves. Despite having 7 times greater available PFD, sun shoots absorbed only 3 times more and had daily carbon gains only double of those of shade shoots. Simulations showed that sun and shade plants performed similarly in the open light environment, but that shade shoots substantially outperformed sun shoots in the shade light environment. The shoot architecture observed in sun plants appears to achieve an efficient compromise between maximizing carbon gain while minimizing the time that the leaf surfaces are exposed to PFDs in excess of those required for light saturation of photosynthesis and therefore potentially photoinhibitory. Received: 8 June 1997 / Accepted: 2 November 1997     

Photographic estimation of photosynthetically active radiation: evaluation of a computerized technique

Summary We describe a computerized technique for the analysis of hemispherical photographs of sites within plant canopies. The analysis yields estimates of canopy openness, a summary of the spatial distribution of canopy openings, estimates of direct and diffuse site factors, predicted times and durations of sunflecks for any date, and predicted daily courses of photosynthetically active photon flux density (PFD) for any date and for horizontal as well as inclined surfaces. By comparing estimates from photographs with measured values in 32 tropical forest microsites, we evaluate this technique and assess its potential and limitations. In sites of widely varying light availability, and with predictions specific to leaves at various angles, measured daily PFD correlated strongly (r=0.908) with predictions from photographs. Measured daily PFD was also closely correlated with estimates, from the photographs, of diffuse and direct site factors (r>0.8). Differences between predictions and measurements were proportionately largest in heavily shaded sites. Under clear conditions, this technique reliably predicted both daily PFD and the temporal pattern of PFD including predicting suflecks lasting more than a few minutes. However, temporal patterns of PFD predicted from photographs differ in potentially important details from measured patterns.C.I.W.D.P.B. Publication no. 935     

Effect of leaf flutter on the light environment of poplars

The dynamics of the canopy light environment for two poplar species (Populus tremuloides Michx., and P. fremontii Wats.) were characterized with an array of photocells in fixed positions within the canopy or attached directly to leaves and using a data logger that recorded photon flux density (PFD) at frequencies from 1 to 20 Hz. The majority of sunflecks were short in duration (<1 s) with a similar short interval between sunflecks. Sunflecks contribute as much as 90% of the total daily PFD in the lower canopy. Leaf flutter may cause high frequency (3 to 5 Hz) variations of PFD in poplar canopies. The amount of light intercepted by a fluttering leaf at the top of the canopy decreased with increasing flutter, whereas a fluttering lower canopy leaf showed no such trend. When leaves fluttered at the top of the canopy the understory light environment showed an increased number of shorter sunflecks. Leaf flutter may increase mean PFD for understory leaves. It also creates a canopy light environment that is more dynamic temporally and more evenly distributed spatially. The potential benefits of these changes in light dynamics are discussed.     

Characteristics of transient photosynthesis in Quercus serrata seedlings grown under lightfleck and constant light regimes

Photosynthetic-induction response and light-fleck utilization were investigated for the current-year seedlings of Quercus serrata, a deciduous tree found in temperate regions of Japan. The tree seedlings were grown under three light regimes: a constant low photosynthetic photon flux density (PFD) regime of 50 mol m^–2 s^–1, a constant high PFD regime of 500 mol m^–2 s^–1, and a lightfleck regime with alternated low (lasting 5 s) and high (lasting 35 s) PFD. The photosynthetic-induction response following a sudden increase of PFD from 50 to 500 mol m^–2 s^–1 exhibited two phases: an initial fast increase complete within 3–5 s, and a second slow increase lasting for 15–20 min. Induction times required to reach 50% and 90% of steady-state assimilation rates were significantly shorter in leaves from the constant low PFD than those from the high PFD regime. During the first 60–100 s, the ratio of observed integrated CO₂ uptake to that predicted by assuming that a steady-state assimilation would be achieved instantaneously after the light increase was significantly higher for leaves from the low PFD regime than from the high PFD regime. Lightfleck utilization was examined for various durations of PFD of 500 mol m^–2 s^–1 on a background PFD of 50 mol m^–2 s^–1. Lightfleck utilization efficiency was significantly higher in low PFD leaves than in the high PFD leaves for 5-s and 10-s lightflecks, but showed no difference among different light regimes for 100-s lightflecks. The contribution of post-illumination CO₂ fixation to total carbon gain decreased markedly with increasing lightfleck durations, but exhibited no significant difference among growth regimes. Photosynthetic performances of induction response and lightfleck utilization in leaves from the lightfleck regime were more similar to those in leaves from the low PFD regime. It may be the total daily PFD rather than PFD dynamics in light regimes that affects the characteristics of transient photosynthesis in Q. serrata seedlings.     

Rapid changes in xanthophyll cycle-dependent energy dissipation and photosystem II efficiency in two vines, Stephania japonica and Smilax australis, growing in the understory of an open Eucalyptus forest

Leaves of Stephania japonica and Smilax australis were characterized in situ on the coast of north-eastern New South Wales, Australia, where they were growing naturally in three different light environments: deep shade, in the understory of an open Eucalyptus forest where they received frequent sunflecks of high intensity, and in an exposed site receiving full sunlight. In deep shade the xanthophyll cycle remained epoxidized during the day and the vast majority of absorbed light was utilized for photosynthesis. In the exposed site both deepoxidation and epoxidation of the xanthophyll cycle and changes in the level of xanthophyll-dependent thermal energy dissipation largely tracked the diurnal changes in photon flux density (PFD). In the understory the xanthophyll cycle became largely deepoxidized to zeaxanthin and antheraxanthin upon exposure of the leaves to the first high intensity sunfleck and this high level of deepoxidation was maintained throughout the day both during and between subsequent sunflecks. In contrast, thermal energy dissipation activity, and the efficiency of photosystem II, fluctuated rapidly in response to the changes in incident PFD. These findings suggest a fine level of control over the engagement of zeaxanthin and antheraxanthin in energy dissipation activity, presumably through rapid changes in thylakoid acidification, such that they became rapidly engaged for photoprotection during the sunflecks and rapidly disengaged upon return to low light when continued engagement might limit carbon gain.