Photosynthetic symmetry of sun and shade leaves of different orientations

Summary The photosynthetic responses to light of leaves irradiated on the adaxial or abaxial surfaces, were measured for plants with contrasting leaf orientations. For vertical-leaf species of open habitats (Eryngium yuccifolium and Silphium terebinthinaceum), photosynthetic rates were identical when irradiated on either surface. However, for horizontal-leaf species of open habitats (Ambrosia trifida and Solidago canadensis), light-saturated rates of photosynthesis for adaxial irradiation were 19 to 37% higher than rates for abaxial irradiation. Leaves of understory plants (Asarum canadense and Hydrophyllum canadense) were functionally symmetrical although they had horizontal orientation. Photosynthetic rates were measured at saturating CO₂, thus differences in the response to incident irradiance presumably resulted from complex interactions of light and leaf optical properties rather than from stomatal effects. Differences in absorptance (400–700 nm) among leaf surfaces were evident for horizontal-leaf species but the primary determinant of functional symmetry was leaf anatomy. Functionally symmetrical leaves had upper and lower palisade layers of equal thickness (vertical leaves of open habitats) or were composed primarily of a single layer of photosynthetic cells (horizontal leaves of understory habitats). Photosynthetic symmetry of vertical-leaf species may be an adaptation to maximize daily integrated carbon gain and water-use efficiency, whereas asymmetry of horizontal-leaf species may be an adaptation to maximize daily integrated carbon gain and photosynthetic nutrient-use efficiency.     

Photosynthetic induction times in shade-tolerant species with long and short-lived leaves

In the understory of a tropical rainforest, light flecks can contribute 10–80% of the total light flux. We investigated the capacity of eight shade-tolerant species to use light flecks by examining the time required for full induction of photosynthesis during an artificial light fleck. CO₂ fixation rates were measured with a portable LiCor gas-exchange system for plants growing in the field on Barro Colorado Island, Panama. In all species induction to 50% of maximum CO₂ fixation occurred quickly, from 1 to 3 min. In species with short leaf lifetimes (1 year), induction to 90% of maximum also occurred quickly, in 3–6 min. In contrast, the species with longer lived leaves (>4 years) required 11–36 min for induction to 90% of maximum. Induction times for leaves from gap and understory plants of the same species were indistinguishable. Elevated CO₂ did not eliminate the slow induction phase of long-lived leaves. This suggests that slow induction did not result from stomatal limitation. O₂ evolution, measured on excised leaf disks, induced in 3–4 min in species with short-lived leaves, and 4–8 min in species with long-lived leaves. The rapid induction of O₂ evolution indicates that the slower induction of CO₂ fixation in long-lived leaves was not caused by a delay in the induction of electron transport. Activation of rubisco may be the major factor limiting response times in species with long-lived leaves. Species from Panama with short-lived leaves had remarkably rapid induction times that are comparable to those of algae or higher plant chloroplasts. We also found that understory forest plants induced two to seven times more quickly than did potted plants.     

四种热带雨林树种光合和形态解剖特征对不同生长光强的适应

研究了西双版纳热带雨林2种喜光树种中平树(Macaranga denticulata)、倒樱木(Pnravallaris macrophylla)和2种耐荫树种云南肉豆蔻(Myristica yunnanensis)、金丝李(Garcinia paucinervis)幼苗叶片光合和形态解剖特征对3种不同生长光强(5％、25%和50％相对光强)的适应。研究结果表明,与强光下相比,弱光下生长的4种植物最大净光合速率、光饱和点、光补偿点、暗呼吸速率、叶绿素a／b、叶片和栅栏组织厚度、气孔密度和比叶重都降低,而海绵组织／栅栏组织和叶绿素含量升高。在相同光强下,与2种耐荫树种相比,2种喜光树种有较大的最大净光合速率、暗呼吸速率、气孔密度和较低的叶绿素含量。在不同光强下,4种植物均表现出了对光适应有利的生理和形态解剖可塑性,而喜光树种比耐荫树种有较大的生理和形态可塑性,表明喜光树种具有比耐荫树种对强光有更强的适应能力。4种植物的生理指标的可塑性均大于叶片解剖结构的可塑性。     

Photosynthetic acclimation in the context of structural constraints to carbon export from leaves

The potential role of foliar carbon export features in the acclimation of photosynthetic capacity to differences and changes in light environment was evaluated. These features included apoplastic vs. symplastic phloem loading, density of loading veins, plasmodesmatal frequency in intermediary cells, and the ratio of loading cells to sieve elements. In initial studies, three apoplastic loaders (spinach, pea, Arabidopsis thaliana) exhibited a completely flexible photosynthetic response to changing light conditions, while two symplastic loaders (pumpkin, Verbascum phoeniceum), although able to adjust to different long-term growth conditions, were more limited in their response when transferred from low (LL) to high (HL) light. This suggested that constraints imposed by the completely physical pathway of sugar export might act as a bottleneck in the export of carbon from LL-acclimated leaves of symplastic loaders. While both symplastic loaders exhibited variable loading vein densities (low in LL and high in HL), none of the three apoplastic loaders initially characterized exhibited such differences. However, an additional apoplastic species (tomato) exhibited similar differences in vein density during continuous growth in different light environments. Furthermore, in contrast to the other apoplastic loaders, photosynthetic acclimation in tomato was not complete following a transfer from LL to HL. This suggests that loading vein density and loading cells per sieve element, and thus apparent loading surface capacity, play a major role in the potential for photosynthetic acclimation to changes in light environment. Photosynthetic acclimation and vein density acclimation were also characterized in the slow-growing, sclerophytic evergreen Monstera deliciosa. This evergreen possessed a lower vein density during growth in LL compared to HL and exhibited a more severely limited potential for photosynthetic acclimation to increases in light environment than the rapidly-growing, mesophytic annuals.     

三种高山杜鹃的光合生理生态研究

对大白花杜鹃（Rhododendron decorum）、云南杜鹃（R.yunnanense）和红棕杜鹃（R.rubiginosum）进行了气体交换、叶片性状等研究,以期了解三种杜鹃的光合生理特性及其对环境的适应。结果表明,三种杜鹃的光饱和光合速率（Pmax）与RuBP饱和最大羧化速率（Vc max）、光饱和最大电子传递速率（Jmax）和气孔导度（gs）呈极显著正相关（P≤0.01）,但仅Vc max存在显著的种间差异,说明三种杜鹃的光合能力主要受Vc max影响。叶氮含量、叶片氮在电子传递和在Rubisco中的分配均显著影响Vc max和Jmax。大白花杜鹃的LSP最低,LCP较高,对强光和弱光利用能力都不强,光适应范围较窄。云南杜鹃LCP最低,LSP和Pmax相对较高,对弱光或较强的光照均能利用,光照适应范围相对最广,光合适应能力最强;红棕杜鹃LSP和LCP均为最高,对强光环境的适应性最强。     

Photosynthetic acclimation in eastern hemlock [Tsuga canadensis (L.) Carr.] seedlings following transfer of shade-grown seedlings to high light

 We studied photosynthetic acclimation of eastern hemlock [Tsuga canadensis (L.) Carr.] seedlings in the first month after sudden exposure of shade-grown seedlings to full sunlight. In a greenhouse experiment, seedlings were grown under full sun or 80% shade, and after 7 months, a sample of the shaded trees was transferred to full sun in the greenhouse. Photosynthetic responses of shaded, transferred, and sun trees were followed over the course of 26 days to track short to medium-term acclimation responses. A partial acclimation of photosynthesis at high light occurred in pre-existing (formed in the previous environment) and new foliage of transferred seedlings. This was associated with non-stomatal limitations to photosynthesis. Pre-existing foliage of transferred plants had a prolonged reduction in the ratio of variable to maximal fluorescence, and a limited capacity to adjust photochemical quenching or photosystem II quantum yield in the light to increasing light intensity compared to sun foliage, and apparently had some difficulty sustaining non-photochemical quenching. Seedling survival was only 58% among transferred seedlings, compared to 80% and 100% in the shade or sun groups, respectively. Photosystem II quantum yield in the light, and photochemical and non-photochemical quenching were similar between newly formed foliage of transferred and sun plants. These findings indicate that eastern hemlock depends strongly on the production of new foliage for photosynthetic adjustments to high light, and that development of photosynthetic competence may be a gradual process that occurs over successive foliar production cycles. Received: 12 May 1998 / Accepted: 27 July 1998     

Photosynthetic light acclimation in fully developed leaves of the juvenile and adult life phases of Hedera helix

The aim of this study was to investigate the extent to which fully developed leaves of Hedera helix L. are capable of acclimating to new light conditions and how this ability is determined by the life phase of the plant. To this end juvenile and adult plants were transferred from a low (L) to a moderately high (H) light regime and vice versa and changes of photosynthetic gas exchange, RuBP carboxylase (EC 4.1.1.39) activity and specific anatomy were monitored in leaves that were fully developed prior to the transfer.
Immediately after transfer from L to H there was a decrease in the rate of net photosynthesis (F_n). This photoinhibition was particularly pronounced in leaves of the adult life phase. F_n recovered after 10 to 20 days at H, and 40 to 65 days after transfer the rate exceeded that of control plants by about 20% in leaves of the adult life phase and by about 50% in leaves of the juvenile life phase. If H plants were transferred to L, F_n had declined only slightly after 30 to 40 days and regained its initial level within a few days, when the plants were returned to the original high light regime.
The increased rates of F_n per unit leaf area in leaves transferred from L to H were associated with higher light levels necessary to saturate F_n, higher carboxylation efficiencies, higher contents of soluble protein and higher activities of RuBP carboxylase, whereas the quantum yield did not change. Although fully differentiated before transfer, the leaves had formed a further cell layer in the palisade parenchyma. Related to leaf volume there was no increase in F_n.
Our results indicate that in the adult life phase of ivy phenotypie light acclimation occurs mainly during leaf development, whereas in juvenile plants fully expanded leaves still possess a rather wide modulativc acclimation plasticity.     

Physiological acclimation to light in Chara intermedia nodes

In this study we investigated the ability of Chara intermedia to acclimate to different irradiances (i.e. “low-light” (LL): 20–30 μmol photons m⁻² s⁻¹ and “high-light” (HL): 180–200 μmol photons m⁻² s⁻¹) and light qualities (white, yellow and green), using morphological, photosynthesis, chlorophyll fluorescence and pigment analysis.Relative growth rates increased with increasing irradiance from 0.016 ± 0.003 (LL) to 0.024 ± 0.005 (HL) g g⁻¹ d⁻¹ fresh weight and were independent of light quality. A growth-based branch orientation towards high-light functioning as a mechanism to protect the plant from excessive light was confirmed. It was shown that the receptor responsible for the morphological reaction is sensitive to blue-light.C. intermedia showed higher oxygen evolution (up to 10.5 (HL) vs. 4.5 (LL) nmol O₂ mg Chl⁻¹ s⁻¹), photochemical and energy-dependent Chl fluorescence quenching and a lower Fv/Fm after acclimation to HL. With respect to qP, the acclimation of the photosynthetic apparatus depended on light quality and needed the blue part of the spectrum for full development. In addition, pigment composition was influenced by light and the Chl a/Car and Antheraxanthin (A) + Zeaxanthin (Z)/Violaxanthin (V) + A + Z (DES) ratios revealed the expected acclimation behaviour in favour of carotenoid protection under HL (i.e. decrease of Chl a/Car from 3.41 ± 0.48 to 2.30 ± 0.35 and increase of DES from 0.39 ± 0.05 to 0.87 ± 0.03), while the Chl a/Chl b ratios were not significantly affected. Furthermore it was shown that morphological light acclimation mechanisms influence the extent of the physiological modifications.     

Photosynthetic light-harvesting function of carotenoids in higher-plant leaves exposed to high light irradiances

Exposure of barley (Hordeum vulgare L.) leaves to strong white light (1500 μmol photons · m⁻² · s⁻¹) decreased the quantum yield of photosynthetic oxygen evolution in green light preferentially absorbed by carotenoids (Φ-510) but not in red light exclusively absorbed by chlorophylls (Φ-650). This phenomenon was observed to be (i) rapidly induced (within a few minutes), (ii) slowly reversible in darkness (within about 1 h), (iii) insensitive to dithiothreitol and (iv) maximally induced by photon flux densities higher than about 1000 μmol · m⁻² · s⁻¹. Determination of the carotenoid composition of the major light-harvesting complex of PSII (LHCII) and analysis of the thylakoid membrane lipid fluidity before and after strong illumination of barley leaves in the presence or the absence of dithiothreitol showed that the light-induced decrease in the Φ-510/Φ-650 ratio did not require the physical detachment of carotenoids from the pigment antennae. Compared to barley plants grown under moderate light and temperature conditions, plants grown in sustained high irradiance at elevated temperature exhibited (i) a lower Φ-510/Φ-650 ratio, (ii) a reduced size of the functional PSII pigment antenna in green light (but not in red light) and (iii) a marked increase in the amount of free carotenoids found in non-denaturing Deriphat-containing electrophoretic gels of thylakoid membranes. Similarly, the Φ-510/Φ-650 ratio of the LHCII-deficient chlorina-f2 barley mutant was very low compared to the wild type. Separation and quantification of the cis/trans carotenoid isomers of barley leaves revealed that strong illumination did not induce pronounced cis-trans isomerization of xanthophylls. Taken together, the data suggest that the efficiency of energy transfer from carotenoids to chlorophylls varies with the light environment both in the short term and in the long term, with excess light energy noticeably inhibiting the photosynthetic light-harvesting function of carotenoids. The photoprotective significance of this carotenoid decoupling from the chlorophyll antennae is discussed. Received: 28 July 1997 / Accepted: 25 October 1997     

Costs and benefits of photosynthetic light acclimation by tree seedlings in response to gap formation

Some shade leaves increase their photosynthetic capacity (P _max) when exposed to a higher irradiance. The increase in P _max is associated with an increase in chloroplast size or number. To accommodate those chloroplasts, plants need to make thick leaves in advance. We studied the cost and benefit of photosynthetic acclimation in mature leaves of a tree species, Kalopanax pictus Nakai, in a cool-temperate deciduous forest. Costs were evaluated as the additional investment in biomass required to make thick leaves, while the benefit was evaluated as an increase in photosynthetic carbon gain. We created gaps by felling canopy trees and examined the photosynthetic responses of mature leaves of the understorey seedlings. In the shade, leaves of K. pictus had vacant spaces that were not filled by chloroplasts in the mesophyll cells facing the intercellular space. When those leaves were exposed to higher irradiance after gap formation, the area of the mesophyll surface covered by chloroplasts increased by 17% and P _max by 27%. This increase in P _max led to an 11% increase in daily carbon gain, which was greater than the amount of biomass additionally invested to construct thicker leaves. We conclude that the capacity of a plant to acclimate to light (photosynthetic acclimation) would contribute to rapid growth in response to gap formation.     

Flexible and reversible responses to different irradiance levels during photosynthetic acclimation of Cypripedium guttatum

Cypripedium guttatum can be found both in open and shady habitats. Photosynthetic acclimation of C. guttatum to different light availabilities was detected using measurements of chlorophyll fluorescence, photosynthesis and leaf traits. When growing under low light conditions, C. guttatum exhibited a greater efficiency in photochemical utilization of absorbed light energy, and a lower ability for non-photochemical dissipation of excess light energy, as compared to the plants growing under high light conditions. Under intermediate light conditions, C. guttatum exhibited higher photosynthetic capacity (A(max)) than those under both low light or high light conditions. The differences in A(max) among three light environments was linked to the differences in biochemical efficiency, leaf N content (LNC) and leaf dry mass per unit area (LMA), but not to the differences of chlorophyll content. However, there were no significant differences in the light compensation points (LCP) and light saturation points (LSP) for photosynthesis for the plants growing under the three light conditions. These results indicate that the photosynthetic capacity of C. guttatum leaves allows for flexible and reversible responses to different irradiance levels. Photosynthetic acclimation in C. guttatum was affected by biochemical changes, the changes in LMA and ratio of Chl a/b. Successful acclimation of C. guttatum to a broad range of light levels likely allows for its wide geographical distribution. A level of about 45% sunlight appears to be optimal for photosynthesis.     

Ecophysiological consequences of non-random leaf orientation in the prairie compass plant,Silphium laciniatum

Summary The prairie compass plant (Silphium laciniatum L.) has vertical leaves that are characteristically oriented in a north-south plane (i.e., the flat surfaces of the lamina face east and west). We explored the consequences of this orientation by determining basic photosynthetic and water use characteristics in response to environmental factors and by determining total daily photosynthesis and water use of leaves held in different orientations. Average maximum CO₂ exchange rate (CER) of leaves near Ames, IA was constant at 22 micromol m^–2 s^–1 from May through August and then declined. CER did not exhibit a distinct lightsaturation point. CER at photon flux densities near full sunlight was constant from 22 to 35°C leaf temperature but declined at higher temperatures. However, leaf temperatures rarely exceed 35°C during the growing season. There was no change in the pattern of response of CER to temperature over the growing season. We constrained leaves to face east-west (EW,=natural), to face north-south (NS), or to be horizontal (HOR) on eight days in 1986–1988. EW leaves had the highest light interception, leaf temperatures, CER, and transpiration early and late in the day, whereas HOR leaves had the highest values in the middle of the day. Integrations of CER and transpiration over the eight daytime periods showed EW and HOR leaves to have equivalent carbon gain, higher than that of NS leaves. HOR leaves had the highest daily transpiration. Daily water use efficiency (WUE, carbon gained/water lost) was always highest in EW leaves, with the HOR leaves having 16% lower WUE and NS leaves having 33% lower WUE. The natural orientation of compass plant leaves results in equivalent or higher carbon gain and in increased WUE when compared to leaves with other possible orientations; this is likely to have a selective advantage in a prairie environment.     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

Fluorescence induction in whole leaves: Differentiation between the two leaf sides and adaptation to different light regimes

In a variety of plants, the induction kinetics of chlorophyll fluorescence vary substantially depending on whether measured on the upper or lower side of the same leaf. The responses are comparable to those of plants grown under sun and shade conditions. Leaf morphology appears not to be the primary cause of the differences since inversion of the leaves can lead to reversed fluorescence responses. Fluorescence induction was analyzed in control and inverted leaves, and in one case, in chloroplasts from sun and shade leaves. It is concluded from the data that the major differences between the chloroplasts of the upper and lower leaf side reflect ionic and thylakoidmembrane conformational factors, rather than structural differences. Mg²⁺ flux probably plays a significant role in the adjustment of the thylakoid membrane to high or low light conditions.     

Photosynthetic light acclimation in two rainforest Piper species with different ecological amplitudes

Summary Piper auritum (H.B. & K.), a pioneer tree restricted to open sites and Piper hispidum (Swartz), a shrub common in sites ranging from recent clearings to shaded understory, both adjust photosynthetic characteristics in response to light availability during growth. The sensitivity of photosynthetic capacity to light availability during growth was indistinguishable for the two species growing in their natural habitat. Photosynthetic capacity was strongly correlated with leaf nitrogen in both species, and the relationship was similar between species. Dark respiration and leaf specific mass were more sensitive to light during growth in P. hispidum, the species with the broad habitat ange, than in P. auritum. In general, similarities between the species were more striking than differences between them. The differences in dark respiration could have important implications for carbon balance. The difference in the responsiveness of leaf specific mass to light indicates that the broad-ranging species maintains access to modes of response little utilized by the open-site specialist. We did not and, in the gas exchange characteristics, any evidence that the open site specialist is better suited than the generalist to high-light sites.This CIW DPB publication number 962     

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.     

Photosynthetic acclimation in shade-developed leaves of Euterpe edulis Mart (Arecaceae) after long-term exposure to high light

To analyze acclimation of Euterpe edulis seedlings to changes in light availability, we transferred three-year-old seedlings cultivated for six months under natural shade understory [≈ 1.3 mol(photon) m^?2 d^?1] to a forest gap [≈ 25.0 mol(photon) m^?2 d^?1]. After the transfer, changes in chlorophyll fluorescence and leaf gas-exchange parameters, as well as in the light-response curves of photosynthesis and photosynthetic induction parameters, were analyzed during the following 110 days. Simultaneously measured photosynthetic characteristics in the shaded seedlings grown in understory served as the control. Despite the fact that the understory seedlings were under suboptimal conditions to achieve their light-saturated net photosynthetic rate (P _Nmax), light-response curves and photosynthetic induction parameters indicated that the species had the low respiration rate and a fast opening of stomata in response to the intermittent occurrence of sunflecks, which exerted a feed-forward stimulation on P _Nmax. Sudden exposure to high light induced photoinhibition during the first week after the transfer of seedlings to gap, as it was shown by the abrupt decline of the maximal quantum yield of PSII photochemistry (F_v/F_m). The photoinhibition showed the time-dependent dynamics, as the F_v/F_m of the seedlings transferred to the forest gap recovered completely after 110 days. Furthermore, the net photosynthetic rate increased 3.5-fold in relation to priorexposure values. In summary, these data indicated that more than 21 days was required for the shade-acclimated seedlings to recover from photoinhibition and to relax induction photosynthetic limitations following the sudden exposure to high light. Moreover, the species responded very quickly to light availability; it highlights the importance of sunflecks to understory seedlings.     

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.     

Photosynthetic acclimation of higher plants to growth in fluctuating light environments

This paper describes a study into the potential of plants to acclimate to light environments that fluctuate over time periods between 15 min and 3 h. Plants of Arabidopsis thaliana (L.) Heynh., Digitalis purpurea L. and Silene dioica (L.) Clairv. were grown at an irradiance 100 mol m^-2 s^-1. After 4–6 weeks, they were transferred to light regimes that fluctuated between 100 and either 475 or 810 mol m^-2 s^-1, in a regular cycle, for 7 days. Plants were shown, in most cases, to be able to undergo photosynthetic acclimation under such conditions, increasing maximum photosynthetic rate. The extent of acclimation varied between species. A more detailed study with S. dioica showed that this acclimation involved changes in both Rubisco protein and cytochrome f content, with only marginal changes in pigment content and composition. Acclimation to fluctuating light, at the protein level, did not fully reflect the acclimation to continuous high light - Rubisco protein increased more than would be expected from the mean irradiance, but less than expected from the high irradiance; cytochrome f increased when neither the mean nor the high irradiance would be expected to induce an increase.This revised version was published online in October 2005 with corrections to the Cover Date.     

Effects of blue light deficiency on acclimation of light energy partitioning in PSII and CO2 assimilation capacity to high irradiance in spinach leaves

Blue light effects on the acclimation of energy partitioningcharacteristics in PSII and CO₂ assimilation capacity in spinachto high growth irradiance were investigated. Plants were grownhydroponically in different light treatments that were a combinationof two light qualities and two irradiances, i.e. white lightand blue-deficient light at photosynthetic photon flux densities(PPFDs) of 100 and 500 µmol m^–2 s^–1. The CO₂assimilation rate, the quantum efficiency of PSII (PSII) andthermal dissipation activity / in young, fully expanded leaves were measured under 1,600 µmol m^–2 s^–1white light. The CO₂ assimilation rate and PSII were higher,while / was lower in plants grown under high irradiancethan in plants grown under low irradiance. These responses wereobserved irrespective of the presence or absence of blue lightduring growth. The extent of the increase in the CO₂ assimilationrate and PSII and the decrease in / by high growth irradiance was smaller under blue light-deficient conditions. These resultsindicate that blue light helps to boost the acclimation responsesof energy partitioning in PSII and CO₂ assimilation to highirradiance. Similarly, leaf N, Cyt f and Chl contents per unitleaf area increased by high growth irradiance, and the extentof the increment in leaf N, Cyt f and Chl was smaller underblue light-deficient conditions. Regression analysis showedthat the differences in energy partitioning in PSII and CO₂assimilation between plants grown under high white light andhigh blue-deficient light were closely related to the differencein leaf N.