Leaf anatomy and light acclimation in woody seedlings after gap formation in a cool-temperate deciduous forest

The photosynthetic light acclimation of fully expanded leaves of tree seedlings in response to gap formation was studied with respect to anatomical and photosynthetic characteristics in a natural cool-temperate deciduous forest. Eight woody species of different functional groups were used; two species each from mid-successional canopy species (Kalopanax pictus and Magnolia obovata), from late-successional canopy species (Quercus crispula and Acer mono), from sub-canopy species (Acer japonicum and Fraxinus lanuginosa) and from vine species (Schizophragma hydrangeoides and Hydrangea petiolaris). The light-saturated rate of photosynthesis (P _max) increased significantly after gap formation in six species other than vine species. Shade leaves of K. pictus, M. obovata and Q. crispula had vacant spaces along cell walls in mesophyll cells, where chloroplasts were absent. The vacant space was filled after the gap formation by increased chloroplast volume, which in turn increased P _max. In two Acer species, an increase in the area of mesophyll cells facing the intercellular space enabled the leaves to increase P _max after maturation. The two vine species did not significantly change their anatomical traits. Although the response and the mechanism of acclimation to light improvement varied from species to species, the increase in the area of chloroplast surface facing the intercellular space per unit leaf area accounted for most of the increase in P _max, demonstrating the importance of leaf anatomy in increasing P _max.     

Physiological and morphological acclimation of shade-grown tree seedlings to late-season canopy gap formation

Because acclimation to canopy gaps may involve coordination of new leaf production with morphological or physiological changes in existing, shade-developed leaves, we examined both new leaf production and photosynthesis of existing leaves on shade-grown seedlings after exposure to a late-season canopy gap. Midway through the summer, we transferred potted, shade-grown seedlings of four co-occurring temperate deciduous tree species representing a range of shade-tolerance categories and leaf production strategies to gaps. Shade-tolerant Acer saccharum was the least responsive to gap conditions. It produced few new, high-light acclimated leaves and increases in photosynthetic rates of shade-developed leaves appeared stomatally limited. Intermediately shade-tolerant Fraxinus americana and Quercus rubra responded most, by producing new leaves and increasing photosynthetic rates of existing shade-developed leaves to levels not significantly different from gap-grown controls within four weeks of gap exposure. Shade-intolerant Liriodendron tulipifera was intermediate in response. In these species, the degree of shoot-level morphological acclimation (new leaf production) and leaf-level physiological acclimation (photosynthetic increases in existing leaves) appear coupled. Mechanisms of acclimation also appear related to intrinsic patterns of nitrogen use and mobilization, the ability to adjust stomatal conductance, and shade tolerance.     

不同光强下生长的亚热带树苗的光合-光响应特性的比较

鼎湖山亚热带森林中的优势树种马尾松、荷木、黧蒴和黄果厚壳桂的幼苗,盆栽于100%、40%和16%的自然光下生长1年.利用氧电极离体测定叶片的光合-光响应曲线,结果表明不同树种及同一树种不同生长光强处理之间差异明显.比较最大净光合速率(Amax)、光补偿点、光饱和点、光抑制点及光能利用效率(QUE)等的变化可见,黄果厚壳桂为耐阴植物,对高光强的光合适应潜力为有限,100%的自然光明显抑制其最大同化力;其余3种植物均为喜光植物,其中马尾松的光合作用对遮阴的适应能力较差,而荷木和黧蒴则相对较强.本文还对植物的光合生理特性与其演替状态的联系作了初步探讨.     

不同光强下生长的亚热带树苗的光合—光响应特性的比较

鼎湖山亚湖热带森林中的优势树种马尾松、荷木、黛蒴和黄果厚壳桂的幼苗,盆栽于１００％、４０％和１６％的自然光下生长１年。利用氧电极离体测定叶片的光合－光响应曲线,结果表明不同树种及同一种树种不同生长光强处理之间差异明显。比较最大净光合速率、光寂偿点、光饱和点、光抑制点及光能利用效率等的变化可见,黄果厚壳桂为耐阴植物,对高光强的光合适应潜力为有限,１００％,自然光明显抑制最大同化力;其余３种植物均为喜     

Contrasting patterns of photosynthetic acclimation and photoinhibition in two evergreen herbs from a winter deciduous forest

The relationship between the microclimate within an Oak-Hickory forest and photosynthetic characters of two resident evergreen herbs with contrasting leaf phenologies was investigated on a monthly basis for 1 full year. Heuchera americana has leaf flushes in the spring and fall, with average leaf life spans of 6–7 months. Hexastylis arifolia produces a single cohort of leaves each spring with a leaf life span of 12–13 months. We predicted that among evergreen plants inhabiting a seasonal habitat, a species for which the frequency of leaf turnover is greater than the frequency of seasonal extremes would have a greater annual range in photosynthetic capacity than a species that only produced a single flush of leaves during the year. Photosynthetic parameters, including apparent quantum yield, maximum photosynthetic capacity (P_max), temperature of maximum photosynthesis, photochemical efficiency of PSII and leaf nitrogen (N) and chlorophyll concentrations, were periodically measured under laboratory conditions in leaves sampled from natural populations of both species. Mature leaves of both species acclimated to changing understory conditions with the mean seasonal differences being significantly greater for Heuchera than for Hexastylis. Area based maximum photosynthetic rates at 25°C were approximately 250% and 100% greater in winter leaves than summer leaves for Heuchera and Hexastylis respectively. Nitrogen concentrations were highest in winter leaves. Chlorophyll concentrations were highest in summer leaves. Low P_max/N values for these species suggest preferential allocation of leaf nitrogen into non-photosynthetic pools and/or light-harvesting function at the expense of photosynthetic enzymes and electron transport components. Despite the increase in photosynthetic capacity, there was evidence of chronic winter photoinhibition in Hexastylis, but not in Heuchera. Among these ecologically similar species, there appears to be a trade-off between the frequency of leaf production and the balance of photosynthetic acclimation and photoinhibition.     

The gap theory in forest dynamics

Since the late 1970s, ecologists interested in forest dynamics have focused their attention to the responses of individuals, populations and communities to “gaps” which are openings created in the forest canopy. This review intended to introduce some collective knowledge on major subjects of the gap theory in forest dynamics, in relation to gap-disturbance regimes, tree regeneration behaviour and community structure.     

Rapid photosynthetic acclimation of Shorea johorensis seedlings after logging disturbance in Central Kalimantan

This study examined the photosynthetic acclimation of pre-existing Shorea johorensis (Dipterocarpaceae) seedlings to the change in conditions that occurs at the time of logging in Central Kalimantan, Indonesia. The hypothesis was that the seedlings would be unable to acclimate beyond partially open conditions after canopy disturbance caused by logging, therefore limiting the potential for regeneration in the most open areas. Bleaching and reductions in the predawn ratio of variable to maximum fluorescence (F _v /F _m) indicated chronic photoinhibition and damage to the previously shade-adapted leaves of seedlings in an area logged 2 weeks earlier. The majority of seedlings in partially open and open environments of an area logged 3 months earlier were already growing fast. Leaves that had developed in the new environment showed only small reductions in predawn F _v /F _m and large increases in the light saturated rate of photosynthesis (A _max) per unit area when compared to shaded seedlings. Leaves in the most open environments had higher but more variable nitrogen concentrations, A _max per unit area and A _max per unit mass when compared to seedlings in partially open environments. Increases in dark respiration were disproportionately large compared to increases in A _max, and may have been the result of increased investment in photoprotective mechanisms. The response of stomatal conductance to the vapour pressure deficit and leaf temperature was examined, but it suggested only a 10% reduction in daily leaf level carbon gain in open environments. The ratio of leaf area to fine root mass was highest in shade-suppressed and newly exposed seedlings, suggesting a potential hydraulic limitation to transpiration during acclimation. However, rainfall during this period was high and leaf water potentials did not differ between disturbed and undisturbed environments. S. johorensis seedlings were capable of significant acclimation to conditions more extreme than partial canopy opening. Low seedling density after logging during the wet season cannot be explained by a limited potential for photosynthetic acclimation. Received: 14 September 1998 / Accepted: 12 August 1999     

The dynamics of photosynthetic acclimation to changes in light quanlity and quality in three Australian rainforest tree species

Photosynthetic acclimation was studied in seedlings of three subtropical rainforest species representing early (Omalanthus populifolius), middle (Duboisia myoporoides) and late (Acmena ingens) successional stages in forest development. Changes in the photosynthetic characteristics of pre-existing leaves were observed following the transfer of plants between deep shade (1–5% of photosynthetically active radiation (PAR), selectively filtered to produce a red/far-red (R/FR) ratio of 0.1) and open glasshouse (60% PAR and a R/FR ratio of 1.1–1.2), and vice versa. The extent and rate of response of the photosynthetic characteristics of each species to changes in light environment were recorded in this simulation of gap formation and canopy closure/overtopping. The light regimes to which plants were exposed produced significant levels of acclimation in all the photosynthetic parameters examined. Following transfer from high to low light, the light-saturated rate of photosynthesis was maintained near pre-transfer levels for 7 days, after which it decreased to levels which closely approximated those in leaves which had developed in low light. The decrease in photosynthetic capacity was associated with lower apparent quantum yields and stomatal conductances. Dark respiration was the parameter most sensitive to changes in light environment, and responded significantly during the first 4–7 days after transfer. Acclimation of photosynthetic capacity to increases in irradiance was significant in two of the three species studied, but was clearly limited in comparison with that of new leaves produced in the high light conditions. This limitation was most pronounced in the early-successional-stage species, O. populifolius. It is likely that structural characteristics of the leaves, imposed at the time of leaf expansion, are largely responsible for the limitations in photosynthetic acclimation to increases in irradiance.     

Photosynthetic and structural acclimation to light direction in vertical leaves of Silphium terebinthinaceum

The azimuth of vertical leaves of Silphium terebinthinaceum profoundly influenced total daily irradiance as well as the proportion of direct versus diffuse light incident on the adaxial and abaxial leaf surface. These differences caused structural and physiological adjustments in leaves that affected photosynthetic performance. Leaves with the adaxial surface facing East received equal daily integrated irradiance on each surface, and these leaves had similar photosynthetic rates when irradiated on either the adaxial or abaxial surface. The adaxial surface of East-facing leaves was also the only surface to receive more direct than diffuse irradiance and this was the only leaf side which had a clearly defined columnar palisade layer. A potential cost of constructing East-facing leaves with symmetrical photosynthetic capcity was a 25% higher specific leaf mass and increased leaf thickness in comparison to asymmetrical South-facing leaves. The adaxial surface of South-facing leaves received approximately three times more daily integrated irradiance than the abaxial surface. When measured at saturating CO₂ and irradiance, these leaves had 42% higher photosynthetic rates when irradiated on the adaxial surface than when irradiated on the abaxial surface. However, there was no difference in photosynthesis for these leaves when irradiated on either surface when measurements were made at ambient CO₂. Stomatal distribution (mean adaxial/abaxial stomatal density = 0.61) was unaffected by leaf orientation. Thus, the potential for high photosynthetic rates of adaxial palisade cells in South-facing leaves at ambient CO₂ concentrations may have been constrained by stomatal limitations to gas exchange. The distribution of soluble protein and chlorophyll within leaves suggests that palisade and spongy mesophyll cells acclimated to their local light environment. The protein/chlorophyll ratio was high in the palisade layers and decreased in the spongy mesophyll cells, presumably corresponding to the attentuation of light as it penetrates leaves. Unlike some species, the chlorophyll a/b ratio and the degree of thylakoid stacking was uniform throughout the thickness of the leaf. It appears that sun-shade acclimation among cell layers of Silphium terebinthinaceum leaves is accomplished without adjustment to the chlorophyll a/b ratio or to thylakoid membrane structure.     

阔叶红松林林隙结构与树种多样性关系研究

对阔叶红松林林隙与林下不同层次树种多样性进行比较,探讨林隙大小及发育阶段与树种多样性间的关系.结果表明,林隙与林下树种多样性存在显著差异（P<0.01）.与林下相比,林隙更新层树种多样性、丰富度和均匀度增大,生态优势度减小（P<0.01）;演替层树种多样性、丰富度和均匀度减小,生态优势度增大（P<0.0001）.林隙中不同层次树种多样性随着林隙大小呈现出相反的变化趋势.长白山阔叶红松林林隙演替层树种多样性和丰富度总体上呈单峰形变化,中等大小林隙（100～250 m²）中树种多样性和丰富度较高,生态优势度较小.而更新层树种多样性和均匀度在≥250 m²和<100 m²的林隙中最大,在200～250 m²的林隙中最小;生态优势度在200～250 m²的林隙中达到最大.更新层和演替层树种对林隙面积大小反应不同,有利于更新层幼苗建立的林隙面积并不是演替层幼树发育和成活的最适面积;随林隙年龄的增加,更新层和演替层树种多样性在不同层次上呈互补关系.     

Simulation of photosynthetic plasticity in response to highly fluctuating light: an empirical model integrating dynamic photosynthetic induction and capacity

An empirical model was developed to simulate photosynthetic responses of leaves to highly fluctuating light, with a special focus on the functional role of photosynthetic induction and capacity. Based on diurnal courses of light as input data, which were recorded at natural plant sites, we applied this model to simulate the corresponding course of net photosynthesis (output data) for leaves of two neotropical tree species. All six model input parameters (leaf-specific) were obtained via measurements of leaf gas exchange. The model was tested for leaves in their natural environments, characterized by frequent light-flecks. We compared measured carbon gains with computed ones, using a standard steady-state and our induction model. Simulation runs with the steady-state model can result in an immense overestimation of the true situation, by 13.4% at open sites [pioneer species Heliocarpus appendiculatus (Turczaninow)] and by 86.5% at low light environments of the understorey [mid to late successional species Billia colombiana (Planchon and Lindley)]. These significant overestimations, particularly in the understorey, are mainly the consequence of neglecting a dynamic photosynthetic induction under fluctuating light conditions. The model presented here resulted in clearly improved predictions; in open and understorey sites the true carbon gain of leaves was computed with a mean error of less than 7%. As most leaves at natural plant sites are exposed to light environments allowing for dynamic rather than steady-state CO₂ assimilation, the significance of such induction models is evident and is discussed in relation to scaling-up from leaf to canopy and to the whole plant indicating a large potential for errors. Received: 3 May 1999 / Accepted: 9 July 1999     

Ecophysiological responses of Fagus sylvatica seedlings to changing light conditions. I. Interactions between photosynthetic acclimation and photoinhibition during simulated canopy gap formation

Natural regeneration of European beech (Fagus sylvatica L.) establishes under shade, but sudden exposure to high irradiance may occur due to openings in the canopy. To elucidate ecophysiological mechanisms associated with survival of European beech seedlings, the gas exchange, chlorophyll concentrations, and chlorophyll a fluorescence parameters of two different beech populations were studied under changing light conditions. Plants were grown both in a growth chamber and at a natural site (one population) where the seedlings were raised in containers placed in understory and in simulated canopy gaps. Upon exposure to high light in the growth chamber, photosynthetic rates of shade-acclimated leaves of seedlings from both populations increased severalfold and then decreased over several days to the rates of the low-light control seedlings. High-light seedlings always had the highest photosynthetic rates. Initial fluorescence displayed a trend opposite that of photosynthesis; it increased over time, and relative fluorescence and half-time rise declined continuously until the end of experiment to very low values. Exposure to high light of shade-acclimated seedlings resulted in a shift in chlorophyll concentrations to levels intermediate between high-light and low-light seedlings. The light treatment effects were statistically greater than population effects; however, seedlings from the Abetone population were found to be more susceptible to changing light conditions than seedlings from Sicily. Reciprocal light treatments on plants growing at the natural site confirmed the results obtained in the growth chamber experiment. Overall, beech seedlings grown in the field appeared to have a fairly large acclimation potential achieved by plasticity in the photosynthetic apparatus. The lack of pronounced acclimation to high light in seedlings grown in the growth chamber was ascribed to a threshold-type relationship between the acclimation capacity and the level of damage. These observations on the limited potential for acclimation to high light in leaves of European beech seedlings which show a clear capability to exploit sunflecks, are discussed in relation to regeneration following canopy gap formation and reinforce the view of the central role of gap formation in forest dynamics. We conclude that small forest gaps (in which sunflecks play a major role) may present a favorable environment for survival and growth of beech because of their limited ability to acclimate to a sudden increase in irradiance and because of the moderate levels of light stress found in small gaps.     

Growth and ecophysiological acclimation of the foliose lichen Lobaria pulmonaria in forests with contrasting light climates

This study aims to assess biomass and area growth of 600 thalli of the old forest lichen, Lobaria pulmonaria, transplanted to three successional boreal forest stands with (1) natural rainfall regime, (2) additional moistening during dry days, and (3) additional moistening with added nutrients. Mean biomass growth during 100 days varied from 8.3% in the dark young spruce forest to 23.1% in the clear-cut area, with the old forest in between (16.0%). Additional moistening did not enhance lichen growth, probably because the transplantation period was wet. Nutrient additions slightly increased area growth compared to artificial water additions only. Growth was determined by a combination of external (forest stand, site factors) and internal factors (chlorophyll content, biomass per area). Transplants acclimated to high light by increasing thickness and chlorophyll a/b-ratio. Some visible bleaching and a strong positive correlation between chlorophyll content per area and lichen growth in clear-cuts suggest some high light-induced chlorophyll degradation. We believe that biomass growth and natural occurrence of L. pulmonaria is controlled by a delicate balance between light availability and desiccation risk, and that the species is confined to old forests due to a physiological trade-off between growth potential and fatal desiccation damage, both of which increase with increasing light. The discrepancy between potential and realized ecological niches is probably caused by a long-term risk to be killed in open habitats by high light during long periods with no rain.     

Photosynthetic acclimation and photoinhibition on exposure to high light in shade-developed leaves of Fagus crenata seedlings

The physiological response of leaves developed in low light (L) on Fagus crenata seedlings exposed to different levels of high light (H: high light, M: medium light) was studied. Measurements were conducted on potted seedlings in the F. crenata forest understory. The seedlings with leaves developed in L were transferred to H (L–H) and M (L–M) in summer. On exposure to high light, the photochemical efficiency of dark-adapted PSII (F_v/F_m) immediately decreased and was followed by a subsequent recovery in both L–H and L–M leaves. The mean value of F_v/F_m in L–H leaves was lower than that in L–M leaves through experiments, indicating that the degree of photoinhibition in L–H leaves was greater than that in L–M leaves. About 1 month after transfer, 37% and 5% of leaves had fallen in L–H and L–M seedlings, respectively. This result also indicated the greater photoinhibition in L–H leaves. Moreover, the photosynthetic capacity (P_Nmax) of L–H leaves decreased. In contrast, the P_Nmax of L–M leaves increased, although the P_Nmax was lower than that of M control leaves. An increase in the xanthophyll cycle pool (VAZ), indicating an increase of the photoprotective function, was found in both L–H and L–M leaves. Especially, the VAZ pool in L–M leaves was higher than that in M leaves by the end of experiments. L–M leaves may avoid photoinhibition effectively by the decrease in excess light with the increase of the P_Nmax or VAZ pool, compared to L–H leaves. Thus, the physiological acclimation on exposure to high light depended on the degree of high light. To achieve successful photosynthetic acclimation with slight photoinhibition, the variation of light intensity before and after exposure to high light would be an important factor because of the difference in excess light.     

Influence of arbuscular mycorrhizal colonization on whole‐plant respiration and thermal acclimation of tropical tree seedlings

Symbiotic arbuscular mycorrhizal fungi (AMF) are ubiquitous in tropical forests. AMF play a role in the forest carbon cycle because they can increase nutrient acquisition and biomass of host plants, but also incur a carbon cost to the plant. Through their interactions with their host plants they have the potential to affect how plants respond to environmental perturbation such as global warming. Our objective was to experimentally determine how plant respiration rates and responses to warmer environment are affected by AMF colonization in seedlings of five tropical tree species at the whole plant level. We evaluated the interaction between AMF colonization and temperature on plant respiration against four possible outcomes; acclimation does or does not occur regardless of AMF, or AMF can increase or decrease respiratory acclimation. Seedlings were inoculated with AMF spores or sterilized inoculum and grown at ambient or elevated nighttime temperature. We measured whole plant and belowground respiration rates, as well as plant growth and biomass allocation. There was an overall increase in whole plant, root, and shoot respiration rate with AMF colonization, whereas temperature acclimation varied among species, showing support for three of the four possible responses. The influence of AMF colonization on growth and allocation also varied among plant species. This study shows that the effect of AMF colonization on acclimation differs among plant species. Given the cosmopolitan nature of AMF and the importance of plant acclimation for predicting climate feedbacks a better understanding of the patterns and mechanisms of acclimation is essential for improving predictions of how climate warming may influence vegetation feedbacks.     

Geographical pattern in photosynthetic light response of Pinus sylvestris in Europe

1. The geographical aspects in photosynthetic light response and stomatal conductance in the shoots of Pinus sylvestris were studied together with structural properties of shoots and needles. Seven stands within the natural distribution area of P. sylvestris in Europe were chosen. CO₂ exchange, irradiance and stomatal conductance ( g_s ) for water vapour were measured and the maximum photosynthetic rate ( P_m ) was determined from the CO₂ exchange measurements.
2. There was a clear pattern in the average values of P_m along the latitudinal gradient. Highest values of P_m were found in the middle parts of the distribution area and they decreased towards both ends of the transect. The highest value was almost twice as high as the lowest one.
3. The between-site variation explained 70% of the total variation in the maximum photosynthetic rate. P_m was not clearly correlated with any single climatic variable or nitrogen concentration in the needles.
4. P_m was closely coupled with stomatal conductance ( r ²=0·74). The differences in P_m and g_s between the sites is likely to reflect adaptation and acclimation to different climates.     

Photosynthetic acclimation to light in juveniles of two cloud forest tree species

The photosynthetic response of juveniles of Decussocarpus rospigliosii, an emergent primary forest species and shade tolerant in its juvenile stages and Alchornea triplinervia, a gap-colonizing species of tropical cloud forest in Venezuela was studied. Daily courses of microenvironmental variables and gas exchange under contrasting light conditions (gap and understory) were carried out in their natural environment and transplanted to different light regimes (shade and sun) in the field. The photosynthetic response and some anatomical characteristics of plants from different treatments were analyzed in the laboratory. Photosynthetic rates were low for both species, and were negative during some diurnal periods, related to the low photosynthetically active radiation levels obtained at both gap (6% of total radiation) and understory (2%). A. triplinervia shows higher rates (1.5–3.0 molm^{-2 -1}) than D. rospigliosii (0.7–1.1 molm^-2s^-1). Both species showed increased photosynthetic rates when grown in gaps. A. triplinervia did not adjust its maximum photosynthetic rates to the prevailing light conditions. In contrast, D. rospigliosii responded to increased light levels. Both species showed low light compensation points when grown under total shade. There was a partial stomatal closure generally during midday in D. rospigliosii. A. triplinervia presented lower leaf conductances, transpiration rates and lesser stomatal control. Some leaf anatomical characteristics, in both species, were affected by variations in the light regime (i.e. increased leaf thickness, leaf specific weight and stomatal density). These results suggest that both species have the ability to respond to variations in their natural light environments, therefore maintaining a favorable carbon balance during the day.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

Leaf photosynthetic light response: a mechanistic model for scaling photosynthesis to leaves and canopies

1. The response of photosynthesis to radiation is an often-studied but poorly understood process, represented empirically in most photosynthesis models. However, in scaling photosynthesis from leaf to canopy, predictions of canopy photosynthesis are very sensitive to parameters describing the response of leaves to Photosynthetic Photon Flux Density (PPFD).
2. In this study, a mechanistic, yet still simple, approach is presented that models the degree of light saturation in leaves explicitly, assuming a heterogeneous environment of PPFD and chlorophyll.
3. Possible mechanisms determining the ratio of chlorophyll to nitrogen are considered, including a direct dependence on PPFD, a mechanism involving the red/far-red ratio of light in the canopy, and an approach based upon maximizing photosynthesis.
4. Comparison of model predictions with two data sets of light, nitrogen and chlorophyll from canopies of Populus and Corylus suggests that the red/far-red mechanism is the most realistic. The data also show that the trees studied do not always optimize their nitrogen partitioning to maximize photosynthetic yield.
5. We then apply the model to the data sets, to predict the shape of light response curves of leaves within canopies and assess the applicability of simple scaling schemes, in which full acclimation of photosynthesis to PPFD justifies the use of big-leaf models. We conclude that, at least for the data used, basic assumptions of such schemes do not hold.     

Leaf orientation and the response of the xanthophyll cycle to incident light

Summary Leaves from two species, Euonymus kiautschovicus and Arctostaphylos uva-ursi, with a variety of different orientations and exposures, were examined in the field with regard to the xanthophyll cycle (the interconversion of three carotenoids in the chloroplast thylakoid membranes). East-, south-, and west-facing leaves of E. kiautschovicus were sampled throughout the day and all exhibited a pronounced and progressive conversion of violaxanthin to zeaxanthin, followed by a reconversion of zeaxanthin to violaxanthin later in the day. Maximal levels of zeaxanthin and minimal levels of violaxanthin were observed at the time when each leaf (orientation) received the maximum incident light, which was in the morning in east-facing, midday in southfacing, and in the afternoon in west-facing leaves. A very slight degree of hysteresis in the removal of zeaxanthin compared to its formation with regard to incident light was observed. Leaves with a broader range of orientations were sampled from A. uva-ursi prior to sunrise and at midday. All of the examined pigments (carotenoids and chlorophylls) increased somewhat per unit leaf area with increasing total daily photon receipt. The sum of the carotenoids involved in the xanthophyll cycle, violaxanthin + antheraxanthin + zeaxanthin, increased more strongly with increasing growth light than any other pigment. In addition, the amounts of zeaxanthin present at midday also increased markedly with increasing total daily photon receipt. The percentage of the xanthophyll cycle that was converted to zeaxanthin (and antheraxanthin) at peak irradiance was very large (approximately 80%) in the leaves of both E. kiautschovicus and A. uva-ursi. The daily changes in the components of the xanthophyll cycle that paralleled the daily changes in incident light in the leaves of E. kiautschovicus, and the increasing levels of the xanthophyll cycle components with total daily photon receipt in the leaves of A. uva-ursi, are both consistent with the involvement of zeaxanthin (i.e. the xanthophyll cycle) in the photoprotection of the photosynthetic apparatus against damage due to excessive light.Abbreviations A antheraxanthin - EPS epoxidation state of the xanthophyll cycle=(V+0.5A)/(V+A+Z) - PFD photon flux density (400–700 nm) - PFD_i photon flux density incident upon the upper leaf surface - T_air air temperature - T_L leaf temperature - V violaxanthin - Z zeaxanthin