Variations in light energy dissipation in <Emphasis Type="Italic">Woodfordia fruticosa</Emphasis> leaves during expansion

Young leaves of tropical trees frequently appear red in color, with the redness disappearing as the leaves mature. During leaf expansion, plants may employ photoprotective mechanisms to cope with high light intensities; however, the variations in anthocyanin contents, nonphotochemical quenching (NPQ), and photorespiration during leaf expansion are poorly understood. Here, we investigated pigment contents, gas exchange, and chlorophyll (Chl) fluorescence in Woodfordia fruticosa leaves during their expansion. Young red leaves had significantly lower Chl content than that of expanding or mature leaves, but they accumulated significantly higher anthocyanins and dissipated more excited light energy through NPQ. As the leaves matured, net photosynthetic rate, total electron flow through PSII, and electron flow for ribulose-1,5-bisphosphate oxygenation gradually increased. Our results provided evidence that photorespiration is of fundamental importance in regulating the photosynthetic electron flow and CO₂ assimilation during leaf expansion.     

Photosynthetic costs and benefits of abaxial versus adaxial anthocyanins in Colocasia esculenta ‘Mojito’

Main conclusion

Anthocyanins in upper (adaxial) leaf tissues provide greater photoprotection than in lower (abaxial) tissues, but also predispose tissues to increased shade acclimation and, consequently, reduced photosynthetic capacity. Abaxial anthocyanins may be a compromise between these costs/benefits. Plants adapted to shaded understory environments often exhibit red/purple anthocyanin pigmentation in lower (abaxial) leaf surfaces, but rarely in upper (adaxial) surfaces. The functional significance of this color pattern in leaves is poorly understood. Here, we test the hypothesis that abaxial anthocyanins protect leaves of understory plants from photo-oxidative stress via light attenuation during periodic exposure to high incident sunlight in the forest understory, without interfering with sunlight capture and photosynthesis during shade conditions. We utilize a cultivar of Colocasia esculenta exhibiting adaxial and abaxial anthocyanin variegation within individual leaves to compare tissues with the following color patterns: green adaxial, green abaxial (GG), green adaxial, red abaxial (GR), red adaxial, green abaxial (RG), and red adaxial, red abaxial (RR). Consistent with a photoprotective function of anthocyanins, tissues exhibited symptoms of increasing photoinhibition in the order (from least to greatest): RR, RG, GR, GG. Anthocyanic tissues also showed symptoms of shade acclimation (higher total chl, lower chl a/b) in the same relative order. Inconsistent with our hypothesis, we did not observe any differences in photosynthetic CO₂ uptake under shade conditions between the tissue types. However, GG and GR had significantly (39 %) higher photosynthesis at saturating irradiance (A _sat) than RG and RR. Because tissue types did not differ in nitrogen content, these patterns likely reflect differences in resource allocation at the tissue level, with greater nitrogen allocated toward energy processing in GG and GR, and energy capture in RG and RR (consistent with relative sun/shade acclimation). We conclude that abaxial anthocyanins are likely advantageous in understory environments because they provide some photoprotection during high-light exposure, but without the cost of decreased A _sat associated with adaxial anthocyanin-induced shade syndrome.     

The anti-photooxidation of anthocyanins-rich leaves of a purple rice cultivar

In the leaf of rice (Oryza sativa L.) cultivar Yunnan purple rice,the anthocyanins with an obvious absorption peak at 530nm were distributed in the cells of upper and lower epidermis,bulliform tissue and bristle. The maximal photosynthetic oxygen evolution rate and chlorophyll content in flag leaves were 28% and 23%,respectively,more than the common green leaf rice cultivar Chijiaoru-anzhan. Higher chlorophyll content is probably one of the physiological adaptations for enhancing light harvesting capacity of the antenna in photosystems in this cyanic leaves species. Upon the photooxidation of leaf segments mediated by methyl viologen in weak light for 3 days,the distinct bleaching of anthocyanins in purple rice was associated with the reduction of scavenging ability to DPPH· free radical ability and the increase in membrane leakage rate. But almost no changes in contents of flavonoids and total phenolics were observed. Chlorophyll fluorescence parameters Fv/Fo,qP and φPSⅡ decreased with the increase in NPQ and DES of xanthophylls cycle after photooxidation treatment. Green rice leaves showed more decrease in DPPH· scavenging rate and more increase in cell membrane leakage rate but showed a trace of anthocyanins during photooxidation. It is sug-gested that anthocyanin may be a beneficial and primary antioxidant in sun cyanic rice leaves against oxidative stress induced by environmental adversity. And photooxidation could induce different changing patterns of anthocyanins between the tested purple and green rice leaves.     

The anti-photooxidation of anthocyanins-rich leaves of a purple rice cultivar

In the leaf of rice (Oryza sativa L.) cultivar Yunnan purple rice, the anthocyanins with an obvious absorption peak at 530nm were distributed in the cells of upper and lower epidermis, bulliform tissue and bristle. The maximal photosynthetic oxygen evolution rate and chlorophyll content in flag leaves were 28% and 23%, respectively, more than the common green leaf rice cultivar Chijiaoruanzhan. Higher chlorophyll content is probably one of the physiological adaptations for enhancing light harvesting capacity of the antenna in photosystems in this cyanic leaves species. Upon the photooxidation of leaf segments mediated by methyl viologen in weak light for 3 days, the distinct bleaching of anthocyanins in purple rice was associated with the reduction of scavenging ability to DPPH · free radical ability and the increase in membrane leakage rate. But almost no changes in contents of flavonoids and total phenolics were observed. Chlorophyll fluorescence parameters Fv/Fo, qP and ϕ_PSII decreased with the increase in NPQ and DES of xanthophylls cycle after photooxidation treatment. Green rice leaves showed more decrease in DPPH · scavenging rate and more increase in cell membrane leakage rate but showed a trace of anthocyanins during photooxidation. It is suggested that anthocyanin may be a beneficial and primary antioxidant in sun cyanic rice leaves against oxidative stress induced by environmental adversity. And photooxidation could induce different changing patterns of anthocyanins between the tested purple and green rice leaves.     

Why leaves turn red in autumn. The role of anthocyanins in senescing leaves of red-osier dogwood

Why the leaves of many woody species accumulate anthocyanins prior to being shed has long puzzled biologists because it is unclear what effects anthocyanins may have on leaf function. Here, we provide evidence for red-osier dogwood (Cornus stolonifera) that anthocyanins form a pigment layer in the palisade mesophyll layer that decreases light capture by chloroplasts. Measurements of leaf absorbance demonstrated that red-senescing leaves absorbed more light of blue-green to orange wavelengths (495-644 nm) compared with yellow-senescing leaves. Using chlorophyll a fluorescence measurements, we observed that maximum photosystem II (PSII) photon yield of red-senescing leaves recovered from a high-light stress treatment, whereas yellow-senescing leaves failed to recover after 6 h of dark adaptation, which suggests photo-oxidative damage. Because no differences were observed in light response curves of effective PSII photon yield for red- and yellow-senescing leaves, differences between red- and yellow-senescing cannot be explained by differences in the capacities for photochemical and non-photochemical light energy dissipation. A role of anthocyanins as screening pigments was explored further by measuring the responses PSII photon yield to blue light, which is preferentially absorbed by anthocyanins, versus red light, which is poorly absorbed. We found that dark-adapted PSII photon yield of red-senescing leaves recovered rapidly following illumination with blue light. However, red light induced a similar, prolonged decrease in PSII photon yield in both red- and yellow-senescing leaves. We suggest that optical masking of chlorophyll by anthocyanins reduces risk of photo-oxidative damage to leaf cells as they senesce, which otherwise may lower the efficiency of nutrient retrieval from senescing autumn leaves.     

The Functional Significance of Black-Pigmented Leaves: Photosynthesis,Photoprotection and Productivity in Ophiopogon planiscapus ‘Nigrescens’

Black pigmented leaves are common among horticultural cultivars, yet are extremely rare across natural plant populations. We hypothesised that black pigmentation would disadvantage a plant by reducing photosynthesis and therefore shoot productivity, but that this trait might also confer protective benefits by shielding chloroplasts against photo-oxidative stress. CO₂ assimilation, chlorophyll a fluorescence, shoot biomass, and pigment concentrations were compared for near isogenic green- and black-leafed Ophiopogon planiscapus ‘Nigrescens’. The black leaves had lower maximum CO₂ assimilation rates, higher light saturation points and higher quantum efficiencies of photosystem II (PSII) than green leaves. Under saturating light, PSII photochemistry was inactivated less and recovered more completely in the black leaves. In full sunlight, green plants branched more abundantly and accumulated shoot biomass quicker than the black plants; in the shade, productivities of the two morphs were comparable. The data indicate a light-screening, photoprotective role of foliar anthocyanins. However, limitations to photosynthetic carbon assimilation are relatively small, insufficient to explain the natural scarcity of black-leafed plants.     

Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species

Juvenile leaves in high-light environments commonly appear red as a result of anthocyanin pigments, which play a photoprotective role during light-sensitive ontogenetic stages. The loss of anthocyanin during leaf development presumably corresponds to a decreased need for photoprotection, as photosynthetic maturation allows leaves to utilize higher light intensities. However, the relationship between photosynthetic development and anthocyanin decline has yet to be quantitatively described. In this study, anthocyanin concentration was measured against photopigment content, lamina thickness, anatomical development, and photosynthetic CO(2) exchange in developing leaves of three deciduous tree species. In all species, anthocyanin disappearance corresponded with development of c. 50% mature photopigment concentrations, c. 80% lamina thickness, and differentiation of the mesophyll into palisade and spongy layers. Photosynthetic gas exchange correlated positively with leaf thickness and chlorophyll content, and negatively with anthocyanin concentration. Species with more rapid photosynthetic maturation lost anthocyanin earliest in development. Chlorophyll a/b ratios increased with leaf age, and were lower than those of acyanic species, consistent with a shading effect of anthocyanin. These results suggest that anthocyanin reassimilation is linked closely with chloroplast and whole-leaf developmental processes, supporting the idea that anthocyanins protect tissues until light processing and carbon fixation have matured to balance energy capture with utilization.     

Photoprotection by foliar anthocyanins mitigates effects of boron toxicity in sweet basil (Ocimum basilicum)

Boron (B) toxicity is an important agricultural problem in arid environments. Excess edaphic B compromises photosynthetic efficiency, limits growth and reduces crop yield. However, some purple-leafed cultivars of sweet basil (Ocimum basilicum) exhibit greater tolerance to high B concentrations than do green-leafed cultivars. We hypothesised that foliar anthocyanins protect basil leaf mesophyll from photo-oxidative stress when chloroplast function is compromised by B toxicity. Purple-leafed ‘Red Rubin’ and green-leafed ‘Tigullio’ cultivars, grown with high or negligible edaphic B, were given a photoinhibitory light treatment. Possible effects of photoabatement by anthocyanins were simulated by superimposing a purple polycarbonate filter on the green leaves. An ameliorative effect of light filtering on photosynthetic quantum yield and on photo-oxidative load was observed in B-stressed plants. In addition, when green protoplasts from both cultivars were treated with B and illuminated through a screen of anthocyanic protoplasts or a polycarbonate film which approximated cyanidin-3-O-glucoside optical properties, the degree of photoinhibition, hydrogen peroxide production, and malondialdehyde content were reduced. The data provide evidence that anthocyanins exert a photoprotective role in purple-leafed basil mesophyll cells, thereby contributing to improved tolerance to high B concentrations.     

Anthocyanin contribution to chlorophyll meter readings and its correction

Leaf chlorophyll content is an important physiological parameter which can serve as an indicator of nutritional status, plant stress or senescence. Signals proportional to the chlorophyll content can be measured non-destructively with instruments detecting leaf transmittance (e.g., SPAD-502) or reflectance (e.g., showing normalized differential vegetation index, NDVI) in red and near infrared spectral regions. The measurements are based on the assumption that only chlorophylls absorb in the examined red regions. However, there is a question whether accumulation of other pigments (e.g., anthocyanins) could in some cases affect the chlorophyll meter readings. To answer this question, we cultivated tomato plants (Solanum lycopersicum L.) for a long time under low light conditions and then exposed them for several weeks (4 h a day) to high sunlight containing the UV-A spectral region. The senescent leaves of these plants evolved a high relative content of anthocyanins and visually revealed a distinct blue color. The SPAD and NDVI data were collected and the spectra of diffusive transmittance and reflectance of the leaves were measured using an integration sphere. The content of anthocyanins and chlorophylls was measured analytically. Our results show that SPAD and NDVI measurement can be significantly affected by the accumulated anthocyanins in the leaves with relatively high anthocyanin content. To describe theoretically this effect of anthocyanins, concepts of a specific absorbance and a leaf spectral polarity were developed. Corrective procedures of the chlorophyll meter readings for the anthocyanin contribution are suggested both for the transmittance and reflectance mode.     

Spectral reflectance indices and pigment functions during leaf ontogenesis in six subtropical landscape plants

Pigment combinations are regulated during leaf ontogenesis. To better understand pigment function, alterations in chlorophyll, carotenoid and anthocyanin concentrations were investigated during different leaf development stages in six subtropical landscape plants, namely Ixora chinensis Lam, Camellia japonica Linn, Eugenia oleina Wight, Mangifera indica L., Osmanthus fragrans Lowr and Saraca dives Pierre. High concentrations of anthocyanin were associated with reduced chlorophyll in juvenile leaves. As leaves developed, the photosynthetic pigments (chlorophyll and carotenoid) of all six species increased while anthocyanin concentration declined. Chlorophyll fluorescence imaging of Φ_PSII (effective quantum yield of PSII) and of NPQ (non-photochemical fluorescence quenching) and determination of electron transport rate-rapid light curve (RLC) showed that maximum ETR (leaf electron transport rate), Φ_PSII and the saturation point in RLC increased during leaf development but declined as they aged. Juvenile leaves displayed higher values of NPQ and Car/Chl ratios than leaves at other developmental stages. Leaf reflectance spectra (400–800 nm) were measured to provide an in vivo non-destructive assessment of pigments in leaves during ontogenesis. Four reflectance indices, related to pigment characters, were compared with data obtained quantitatively from biochemical analysis. The results showed that the ARI (anthocyanin reflectance index) was linearly correlated to anthocyanin concentration in juvenile leaves, while a positive correlation of Chl NDI (chlorophyll normalized difference vegetation index) to chlorophyll a concentration was species dependent. Photosynthetic reflectance index was not closely related to Car/Chl ratio, while a structural-independent pigment index was not greatly altered by leaf development or species. Accordingly, it is suggested that the high concentration of anthocyanin, higher NPQ and Car/Chl ratio in juvenile leaves are important functional responses to cope with high radiation when the photosynthetic apparatus is not fully developed. Another two leaf reflectance indices, ARI and Chl NDI, are valuable for in vivo pigment evaluation during leaf development.     

Profiles of photosynthesis within red and green leaves of Quintinia serrata

We have measured photosynthesis at the cellular, tissue, and whole leaf levels to understand the role of anthocyanin pigments on patterns of light utilization. Profiles of chlorophyll fluorescence through sections of red and green leaves of Quintinia serrata showed that anthocyanins in the mesophyll restricted absorption of green light to the uppermost palisade mesophyll. The distribution was further restricted when anthocyanins were also present in the upper epidermis. Mesophyll cells located beneath a cyanic light-filter assumed the characteristic photosynthetic features of shade-adapted cells. As a result, red leaves showed a 23% reduction in CO₂ assimilation under light-saturating conditions, and a lower threshold irradiance for light-saturation, relative to those of green leaves. The photosynthetic characteristics of red leaves are comparable to those of shade-acclimated plants.     

ANATOMICAL AND PHYSIOLOGICAL ACCLIMATION OF FATSIA JAPONICA LEAVES TO IRRADIANCE

Anatomical and physiological leaf characteristics and biomass production of Fatsia japonica plants were studied. Plants were grown in a growth chamber at 300 μmol m^-2 s^-1 (high light) and 50 μmol m^-2 s^-1 (low light) photosynthetic photon flux density. Plants grown under high light showed a net maximum photosynthetic rate 44% higher than plants grown under low light; the light compensation point and the light saturation point were also higher in high-light plants. Photosynthetic oxygen evolution in isolated chloroplasts was about 40% higher in high-light plants. However, chlorophyll content on a dry weight basis, on a leaf area basis, and per chloroplast was greater in plants grown under low light. Leaf thickness in high-light plants was 13% higher than in low-light plants. The number of chloroplasts was 30% higher in high-light leaves, while chloroplast size was only slightly higher. Chloroplast ultrastructure was also affected by light. Leaf dry weight, leaf area, and biomass production per plant were drastically reduced under low light. Thus, F. japonica is a plant that is able to acclimate to different photosynthetic photon flux density by altering its anatomical and physiological characteristics. However, low-light acclimation of this plant has a considerable limiting effect on biomass production.     

Contributions of leaf photosynthetic capacity, leaf angle and self-shading to the maximization of net photosynthesis in Acer saccharum: a modelling assessment

Background and Aims

Plants are expected to maximize their net photosynthetic gains and efficiently use available resources, but the fundamental principles governing trade-offs in suites of traits related to resource-use optimization remain uncertain. This study investigated whether Acer saccharum (sugar maple) saplings could maximize their net photosynthetic gains through a combination of crown structure and foliar characteristics that let all leaves maximize their photosynthetic light-use efficiency (ɛ).

Methods

A functional–structural model, LIGNUM, was used to simulate individuals of different leaf area index (LAI_ind) together with a genetic algorithm to find distributions of leaf angle (L_A) and leaf photosynthetic capacity (A_max) that maximized net carbon gain at the whole-plant level. Saplings grown in either the open or in a forest gap were simulated with A_max either unconstrained or constrained to an upper value consistent with reported values for A_max in A. saccharum.

Key Results

It was found that total net photosynthetic gain was highest when whole-plant PPFD absorption and leaf ɛ were simultaneously maximized. Maximization of ɛ required simultaneous adjustments in L_A and A_max along gradients of PPFD in the plants. When A_max was constrained to a maximum, plants growing in the open maximized their PPFD absorption but not ɛ because PPFD incident on leaves was higher than the PPFD at which ɛ_max was attainable. Average leaf ɛ in constrained plants nonetheless improved with increasing LAI_ind because of an increase in self-shading.

Conclusions

It is concluded that there are selective pressures for plants to simultaneously maximize both PPFD absorption at the scale of the whole individual and ɛ at the scale of leaves, which requires a highly integrated response between L_A, A_max and LAI_ind. The results also suggest that to maximize ɛ plants have evolved mechanisms that co-ordinate the L_A and A_max of individual leaves with PPFD availability.     

Functional responses of Acer species to two simulated forest gap environments: leaf-level properties and photosynthesis

Seedlings of eight forest maple (Acer L.) species were grown outdoors through a full season under two irradiation treatments: (a) “gap edge” with a photosynthetic photon flux density of 30 μmol m-² s-¹ and a red:far-red ratio of 0.55, and (b) “gap centre” with 400 μmol m-² s-¹ and a red:far-red ratio of 1.12. Area-based leaf nitrogen concentration was greater in gap centre-grown seedlings, whereas, except for A. saccharum, area-based chlorophyll (Chl) (a+b) was higher in gap edge-grown plants. There was also a significantly lower Chl a/b ratio in gap edge-grown plants. Maximum photosynthetic rate (P _max ) was 60 % higher in the gap-centre treatment. These results are consistent with the functional expectation that shade-acclimated plants will increase their radiant-energy harvesting capacity as a result of limited photon input while gap-acclimated plants will operate more efficiently under bright irradiance by increasing their carboxylation capacity. This inverse relationship between the capacity of the light-harvesting component and the carboxylation component is, however, only partially supported by Chl fluorescence measurements of intact leaves. Compared to gap centre-grown plants, the lower total fluorescence quenching in gap edge-grown plants indicated a lower carboxylation capacity that was in accord with the observed P _max . However, edge-grown seedlings did not show the expected improvement in light-harvesting efficiency and reduction in electron transport of photosystem 2 inferred from their marginally greater t_1/2 and lower F_v/F_m, respectively. Hence while maples acclimated to different irradiation levels by adjusting leaf N and Chl contents, they showed limited acclimation potential at the photosystem level. Variations in the leaf traits examined had only minor effect on low irradiance photosynthesis and sunfleck utilization. This revised version was published online in August 2006 with corrections to the Cover Date.     

Adaptations of the Photosynthetic Mechanism of Sugar Maple (Acer saccharum) Seedlings Grown in Various Light Intensities

Sugar maple (Acer saccharum Marsh.) seedlings were grown in a nursery for three years in 13, 25, 45 and 100 per cent of full daylight. During the third year of growth, the rates of their apparent photosynthesis and respiration were measured periodically with an infra-red gas analyzer at various light intensities and normal CO₂ concentration. In addition, the rates of apparent photosynthesis of a single attached leaf of the same seedlings were measured at saturating light intensity, hut varying CO₂ concentrations. An increase in the light intensity in which seedlings were grown had no effect on their height or mean leaf area, hut resulted in thicker leaves, an increase in the total leaf area per seedling due to an increase in the number of leaves, an increase in the dry weight especially of roots and a decrease in the chlorophyll content of leaves. Throughout the growing season seedlings grown in full daylight, as compared with those grown in lower light intensities, had the lowest rates of apparent photosynthesis measured at standard conditions (21,600 lux light intensity and 300 ul/l of CO₂), when this was expressed per unit leaf area, hut the highest rates on a per seedling basis. Thus dry matter production attained at the end of the growing season correlated positively with the photosynthetic rate per seedling, but not per unit leaf area. The rates of apparent photosynthesis of seedlings grown at lower light intensities were more responsive to changes in light intensity or CO₂ concentration than those of seedlings grown in full daylight intensity.     

Association of red coloration with senescence of sugar maple leaves in autumn

We evaluated the association of red coloration with senescence in sugar maple (Acer saccharum Marsh.) leaves by assessing differences in leaf retention strength and the progression of the abscission layer through the vascular bundle of green, yellow, and red leaves of 14 mature open-grown trees in October 2002. Computer image analysis confirmed visual categorization of leaves as predominantly green, yellow or red, and chemical quantification of leaf pigment concentrations verified that leaf color reflected underlying differences in leaf biochemistry. Significantly lower chlorophyll concentrations within red and yellow leaves indicated that senescence was more advanced in leaves from these color categories relative to green leaves. Among leaf types, only red leaves contained high concentrations of anthocyanins. There were significant differences in leaf retention capacity among color categories, with the petioles of green leaves being the most firmly attached to twigs, followed by red and then yellow leaves. Microscopic analysis indicated that yellow leaves had the most advanced extension of the abscission layer through the vasculature, with green and red leaves having significantly less abscission layer progression than yellow. A more limited progression of the abscission layer through vascular bundles may be evidence of delayed leaf senescence that could extend resorption of mobile leaf constituents. Together, results from this study suggest an association between leaf anthocyanin content and functional delays in senescence.     

Characteristics of gas exchange and chlorophyll fluorescence in red and green leaves of Begonia semperflorens

To determine the effects of leaf colour on gas exchange and chlorophyll fluorescence, two genotypes of Begonia semperflorens with green leaves or red leaves were compared. The red leaves showed a high accumulation of anthocyanins and high absorbance at 282 and 537 nm while the green leaves exhibited a higher net photosynthetic rate and lower thermal dissipation of light energy. It seems likely that anthocyanins in the vacuoles restricted the absorption of green light to the chloroplasts, leading to a decrease in the efficiency of excitation capture by open PS 2 centres, photochemical quenching and CO₂ assimilation.     

Singlet oxygen and photo-oxidative stress management in plants and algae

Photosynthetic organisms constantly face the threat of photo-oxidative stress from fluctuating light conditions and environmental stress. Plants and algae have developed an array of defences to protect the chloroplast from reactive oxygen species. Genetic and physiological studies have shown that antioxidant responses are important to high-light acclimation, both by directly scavenging or quenching reactive oxygen intermediates and by contributing reducing power for alternative electron transport pathways and excess energy dissipation. At present, the signalling events leading to up-regulation of antioxidant defences in high light remain a mystery. Recent advances toward understanding acclimation to oxidative stress in both photosynthetic and non-photosynthetic model organisms may illuminate how plants and algae respond to high-light stress. Although the role of hydrogen peroxide in high-light acclimation has been investigated, less is known about responses to singlet oxygen, a form of reactive oxygen that poses a significant threat specifically to photosynthetic organisms. This review will discuss some intriguing new findings in that area, focusing on recent findings regarding the nature of singlet-oxygen responses in the chloroplast.     

Effects of moderate high-temperature stress on photosynthesis in three saplings of the constructive tree species of subtropical forest

During the exposition to moderate high-temperature stress, photosynthetic rates and fluorescence of chlorophyll a were measured with a photosynthetic measurement system (Li-Cor 6400) and leaf chamber fluorometer (Li-Cor6400 LCF), respectively, in leaves of saplings, sun-adapted species (Schima superba), shade-adapted species (Cryptocarya concinna), and in mesophytic plant (Castanopsis hystrix) (42°C). The results showed that moderate high-temperature stress led to a decrease in F_v/F_m, namely the primary photochemical quantum efficiency, indicating that moderate high-temperature stress causes a partial inhibition of PSII. It also showed that such an effect was more severe in the shade-adapted plant C. concinna than in the sun-adapted species S. superba. However, except for the sun-grown leaves of C. concinna, the moderate high-temperature stress increased the photosynthetic rate of leaves at high light intensity. Simultaneously, less photoinhibition was found to occur under high-light intensity, suggesting that the capacity of resistant-photoinhibition was stimulated by moderate high-temperature stress. The quantum yield of PSII (ϕ_PSII) decreased in the sun-grown leaves of S. superba and C. hystrix but did not show any significant change in leaves of the shade plant C. concinna and shade-grown leaves of sun plant S. superba or the mesophytic plant C. hystrix because they already had a very low ϕ_PSII under this condition. Moderate high-temperature stress led to a decrease in ϕ_PSII/ϕ_CO2 ratios, an estimate of the quantum requirement for CO₂ assimilation, in the sun plant S. superba and the mesophytic plant C. hystrix because they were associated with the dissipation of a lower fraction of excitation energy. However, no significant changes were found in shade plant C. concinna and in shade-grown leaves of the other examined plants. The effect of moderate high-temperature (42°C) on photosynthesis depends on species and leaf type (sun and shade leaves) in the saplings of subtropical broad-leaved trees.     

Photosynthetic activity,chloroplast ultrastructure,and leaf characteristics of high-light and low-light plants and of sun and shade leaves

The photosynthetic CO₂-fixation rates, chlorophyll content, chloroplast ultrastructure and other leaf characteristics (e.g. variable fluorescence, stomata density, soluble carbohydrate content) were studied in a comparative way in sun and shade leaves of beech (Fagus sylvatica) and in high-light and low-light seedlings.

1. Sun leaves of the beech possess a smaller leaf area, higher dry weight, lower water content, higher stomata density, higher chlorophyll a/b ratios and are thicker than the shade leaves. Sun leaves on the average contain more chlorophyll in a leaf area unit; the shade leaf exhibits more chlorophyll on a dry weight basis. Sun leaves show higher rates for dark respiration and a higher light saturation of photosynthetic CO₂-fixation. Above 2000 lux they are more efficient in photosynthetic quantum conversion than the shade leaves.
2. The development of HL-radish plants proceeds much faster than that of LL-plants. The cotyledons of HL-plants show a higher dry weight, lower water content, a higher ratio of chlorophyll a/b and a higher gross photosynthesis rate than the cotyledons of the LL-plants, which possess a higher chlorophyll content per dry weight basis. The large area of the HL-cotyledon on the one hand, as well as the higher stomata density and the higher respiration rate in the LL-cotyledon on the other hand, are not in agreement with the characteristics of sun and shade leaves respectively.
3. The development, growth and wilting of wheat leaves and the appearance of the following leaves (leaf succession) is much faster at high quanta fluence rates than in weak light. The chlorophyll content is higher in the HL-leaf per unit leaf area and in the LL-leaf per g dry weight. There are no differences in the stomata density and leaf area between the HL- and LL-leaf. There are fewer differences between HL- and LL-leaves than in beech or radish leaves.
4. The chloroplast ultrastructure of shade-type chloroplasts (shade leaves, LL-leaves) is not only characterized by a much higher number of thylakoids per granum and a higher stacking degree of thylakoids, but also by broader grana than in sun-type chloroplasts (sun leaves, HL-leaves). The chloroplasts of sun leaves and of HL-leaves exhibit large starch grains.
5. Shade leaves and LL-leaves exhibit a higher maximum chlorophyll fluorescence and it takes more time for the fluorescence to decline to the steady state than in sun and HL-leaves. The variable fluorescence VF (ratio of fluorescence decrease to steady state fluorescence) is always higher in the sun and HL-leaf of the same physiological stage (maximum chlorophyll content of the leaf) than in the shade and LL-leaf. The fluorescence emission spectra of sun and HL-leaves show a higher proportion of chlorophyli fluorescence in the second emission maximum F₂ than shade and LL-leaves.
6. The level of soluble carbohydrates (reducing sugars) is significantly higher in sun and HL-leaves than in shade and LL-leaves and even reflects changes in the amounts of the daily incident light.
7. Some but not all characteristics of mature sun and shade leaves are found in HL- and LL-leaves of seedlings. Leaf thickness, dry weight, chlorophyll content, soluble carbohydrate level, photosynthetic CO₂-fixation, height and width of grana stacks and starch content, are good parameters to describe the differences between LL- and HL-leaves; with some reservations concerning age and physiological stage of leaf, a/b ratios, chlorophyll content per leaf area unit and the variable fluorescence are also suitable.