Interactive effects of mycorrhization and elevated atmospheric CO2 on sulphur nutrition of young pedunculate oak (Quercus robur L.) trees

Pedunculate oak (Quercus robur L.) was germinated and grown at ambient CO₂ concentration and 650 μmol mol^?1 CO₂ in the presence and absence of the ectomycorrhizal fungus Laccaria laccata for a total of 22 weeks under nonlimiting nutrient conditions. Sulphate uptake, xylem loading and exudation were analysed in excised roots. Despite a relatively high affinity for sulphate (K_M= 1.6 mmol m^?3), the rates of sulphate uptake by excised lateral roots of mycorrhizal oak trees were low as compared to herbaceous plants. Rates of sulphate uptake were similar in mycorrhizal and non-mycorrhizal roots and were not affected by growth of the trees at elevated CO₂. However, the total uptake of sulphate per plant was enhanced by elevated CO₂ and further enhanced by elevated CO₂ and mycorrhization. Sulphate uptake seemed to be closely correlated with biomass accumulation under the conditions applied. The percentage of the sulphate taken up by mycorrhizal oak roots that was loaded into the xylem was an order of magnitude lower than previously observed for herbaceous plants. The rate of xylem loading was enhanced by mycorrhization and, in roots of mycorrhizal trees only, by growth at elevated CO₂. On a whole-plant basis this increase in xylem loading could only partially be explained by the increased growth of the trees. Elevated CO₂ and mycorrhization appeared to increase greatly the sulphate supply of the shoot at the level of xylem loading. For all treatments, calculated rates of sulphate exudation were significantly lower than the corresponding rates of xylem loading of sulphate. Radiolabelled sulphate loaded into the xylem therefore seems to be readily diluted by unlabelled sulphate during xylem transport. Allocation of reduced sulphur from oak leaves was studied by flap-feeding radiolabelled GSH to mature oak leaves. The rate of export of radioactivity from the fed leaves was 4–5 times higher in mycorrhizal oak trees grown at elevated CO₂ than in those grown at ambient CO₂. Export of radiolabel proceeded almost exclusively in a basipetal direction to the roots. From these experiments it can be concluded that, in mycorrhizal oak trees grown at elevated CO₂, the transport of sulphate to the shoot is increased at the level of xylem loading to enable increased sulphate reduction in the leaves. Increased sulphate reduction seems to be required for the enhanced allocation of reduced sulphur to the roots which is observed in trees grown at elevated CO₂. These changes in sulphate and reduced sulphur allocation may be a prerequisite for the positive effect of elevated CO₂ on growth of oak trees previously observed.     

Leaf life span of floating-leaved plants

Photosynthetic capacity of floating-leaved plants is relatively high comparable with terrestrial herbaceous plants, though floating-leaved plants have a much smaller biomass with a leaf area index seldom exceeding 2m²m^-2. Their rather small biomass accumulation is related to higher turnover of leaf biomass or shorter leaf life span. Life span of floating leaves reported in the literature ranged mostly from 13 to 35 days, shorter than that of any other groups of herbaceous macrophytes. Floating-leaved plants are known to show considerably high plasticity in their leaf form. Leaf life span could be prolonged for Nymphoides peltata (Gmel.) O. Kuntze grown in a terrestrial environment and for emergent leaves of Nelumbo nucifera Gaertn. Their short leaf life span seems to be closely related to the fact that old leaves covered by newly formed ones are inevitably compelled to be submerged and lose their function as a photosynthetic apparatus.Abbreviations LAI leaf area index - PFD photosynthetic photon flux density     

Interactive effects of elevated atmospheric CO2, mycorrhization and drought on long-distance transport of reduced sulphur in young pedunculate oak trees (Quercus robur L.)

Pedunculate oak (Quercus robur L.) was germinated and grown under nutrient non-limiting conditions for a total of 10–15 weeks at ambient CO₂ concentration and 1100 μmol mol^–1 CO₂ either in the presence or the absence of the mycorrhizal fungus Laccaria laccata. Half of the oak trees of these treatments were exposed to drought during final growth by suspending the water supply for 21 d. Mycorrhization and elevated atmospheric CO₂ each enhanced total plant biomass per tree. Whereas additional biomass accumulation of trees grown under elevated CO₂ was mainly attributed to increased growth of lateral roots, mycorrhization promoted shoot growth. Water deficiency reduced biomass accumulation without affecting relative water content, but this effect was more pronounced in mycorrhizal as compared to non-mycorrhizal trees. Elevated CO₂ partially prevented the development of drought stress, as indicated by leaf water potential, but did not counteract the negative effects of water deficiency on growth during the time studied. Enhanced biomass accumulation requires adaption in protein synthesis and, as a consequence, enhanced allocation of reduced sulphur produced in the leaves to growing tissues. Therefore, allocation of reduced sulphur from oak leaves was studied by flap-feeding radiolabelled GSH, the main long-distance transport form of reduced sulphur, to mature oak leaves. Export of radiolabel proceeded almost exclusively in basipetal direction to the roots. The rate of export of radioactivity out of the fed leaves was significantly enhanced under elevated CO₂, irrespective of mycorrhization. A higher proportion of the exported GSH was transported to the roots than to basipetal stem sections under elevated CO₂ as compared to ambient CO₂. Mycorrhization did not affect ³⁵S export out of the fed leaves, but the distribution of radiolabel between stem and roots was altered in preference of the stem. Trees exposed to drought did not show appreciable export of the ³⁵S radioactivity fed to the leaves when grown under ambient CO₂. Apparently, drought inhibited basipetal transport of reduced sulphur at the level of phloem loading and/or phloem transport. Elevated CO₂ seemed to counteract this effect of drought stress to some extent, since higher leaf water potentials and improved ³⁵S export out of the fed leaves was observed in oak trees exposed to drought and elevated CO₂ as compared to trees exposed to drought and ambient CO₂.     

Growth and biomass turnover ofHydrocharis dubia L. cultured under different nutrient conditions

Growth of a floating-leaved plant,Hydrocharis dubia L., was examined under varying nutrient conditions between 0.3 and 30 mgN l⁻¹ total inorganic nitrogen.H. dubia plants cultured under the most nutrient-rich condition showed the highest maximum ramet density (736 m⁻²), the highest maximum biomass (80.4 g dry weight m⁻²), and the highest total net production (185 g dry weight m⁻² in 82 days). Plants under nutrient-poor conditions had a relatively large proportion of root biomass and a small proportion of leaves with a long life span. Compared with other floating-leaved and terrestrial plants, the maximum biomass ofH. dubia was relatively small. This, and the rapid biomass turnover, was related to the short life span of leaves (13.2–18.7 days) and large biomass distribution to leaves.     

Sinks for photosynthetic electron flow in green petioles and pedicels of Zantedeschia aethiopica: evidence for innately high photorespiration and cyclic electron flow rates

A combination of gas exchange and various chlorophyll fluorescence measurements under varying O₂ and CO₂ partial pressures were used to characterize photosynthesis in green, stomata-bearing petioles of Zantedeschia aethiopica (calla lily) while corresponding leaves served as controls. Compared to leaves, petioles displayed considerably lower CO₂ assimilation rates, limited by both stomatal and mesophyll components. Further analysis of mesophyll limitations indicated lower carboxylating efficiencies and insufficient RuBP regeneration but almost similar rates of linear electron transport. Accordingly, higher oxygenation/carboxylation ratios were assumed for petioles and confirmed by experiments under non-photorespiratory conditions. Higher photorespiration rates in petioles were accompanied by higher cyclic electron flow around PSI, the latter being possibly linked to limitations in electron transport from intermediate electron carriers to end acceptors and low contents of PSI. Based on chlorophyll fluorescence methods, similar conclusions can be drawn for green pedicels, although gas exchange in these organs could not be applied due to their bulky size. Since our test plants were not subjected to stress we argue that higher photorespiration and cyclic electron flow rates are innate attributes of photosynthesis in stalks of calla lily. Active nitrogen metabolism may be inferred, while increased cyclic electron flow may provide the additional ATP required for the enhanced photorespiratory activity in petiole and pedicel chloroplasts and/or the decarboxylation of malate ascending from roots.     

Physiological and Biochemical Characteristics of Sugar Beet Plants Grown at an Increased Carbon Dioxide Concentration and at Various Nitrate Doses

Three-week-old sugar beet (Beta vulgaris L.) seedlings were grown for an additional four weeks under controlled conditions: in river sand watered with a modified Knop mixture containing one half-fold (0.5N), standard (1N), and or threefold (3N) nitrate amount, at the irradiance of 90 W/m² PAR, and at the carbon dioxide concentrations of 0.035% (1C treatment) or 0.07% (2C treatment). The increase in the carbon dioxide concentration and in the nitrogen dose resulted in an increase in the leaf area and the leaf and root dry weight per plant. With the increase in the nitrogen dose, morphological indices characterizing leaf growth increased more noticeably in 1C plants than in 2C plants. And vice versa, the effects of increased CO₂ concentration were reduced with the increase in the nitrogen dose. Roots responded to the changes in the CO₂ and nitrate concentrations otherwise than leaves. At a standard nitrate dose (1N), the contents of proteins and nonstructural carbohydrates (sucrose and starch) in leaves depended little on the CO₂ concentration. At a double CO₂ concentration, the content of chlorophyll somewhat decreased, and the net photosynthesis rate (P _n) calculated per leaf area unit increased. An increase in the nitrogen dose did not affect the leaf carbohydrate content of the 1C and 2C plants except the leaves of the 2C-3N plants, where the carbohydrate content decreased. In 1C and 2C plants, an increase in the nitrogen dose caused an increase in the protein and chlorophyll content. Specific P _n values somewhat decreased in 1C-0.5N plants and had hardly any dependence on the nitrate dose in the 2C plants. The carbohydrate content in roots did not depend on the CO₂ concentration, and the content was the highest at 0.5N. Characteristic nitrogen dose-independent acclimation of photosynthesis to an increased carbon dioxide concentration, which was postulated previously [1], was not observed in our experiments with sugar beet grown at doubled carbon dioxide concentration.     

Detoxification of SO2 derivatives in chloroplasts ofHardwickia binata

Intact chloroplasts isolated from sulphur dioxide fumigatedHardwickia binata leaves showed inhibition of PS II electron transport activity without any significant effect on photosystem I. Sulphur dioxide exposed leaves accumulated more hydrogen peroxide than those from non-fumigated plants and this was caused by increase in superoxide radical production. Hydrogen peroxide formation was inhibited by addition of cytochrome C and superoxide disrnutase. In sulphur dioxide fumigated leaves, increase in superoxide dismutase activity showed resistance to sulphite toxicity. The localization of ascorbate peroxidase, glutathione reductase and dehydroascorbate reductase activities in chloroplasts provide evidence for the photogeneration of ascorbate. The scavenging of hydrogen peroxide in chloroplast due to ascorbate regenerated from DHA by the system: PS I → Fd → NADP → glutathione. The system can be considered as a means for preliminary detoxification of sulphur dioxide by chloroplasts     

Stomatal and photosynthetic responses ofCichorium intybus leaves to sulfur dioxide treatment at different stages of plant development

Fifty-day-oldCichorium intybus Linn, plants were exposed to 1 ppm sulfur dioxide gas, 2 h per day for 7 consecutive days. Their leaves as well as those from the control plants were sampled at pre-flowering, flowering, and post-flowering stages to study their morphological, physiological, and biochemical responses to SO₂ stress. The number, dimensions, area, and biomass of leaves were less in the treated plants. Length and width of stomatal apertures on both epidermises were greater for leaves exposed to SO₂. The Stomata were longer on the adaxial epidermis, but shorter on the abaxial epidermis, except at the pre-flowering stage. Stomatal widths varied widely. Compared with the controls, the abaxial epidermis on treated leaves showed consistently lower stomatal densities as well as stomatal indices. This was also true for the adaxial epidermis during the post-flowering stage. The photosynthetic rate and stomatal conductance were reduced in the SO₂-exposed plants, but intercellular CO₂ concentrations increased at the pre-flowering stage and, subsequently, declined. Chlorophyll a, carotenoid, and total chlorophyll contents increased at the pre-flowering stage, and then decreased. The level of chlorophyllb was reduced throughout plant development compared with the untreated controls.     

Sagebrush and grasshopper responses to atmospheric carbon dioxide concentration

Summary Seed- and clonally-propagated plants of Big Sagebrush (Artemisia tridentata var.tridentata) were grown under atmospheric carbon dioxide regimes of 270, 350 and 650 μl l⁻¹ and fed toMelanoplus differentialis andM. sanguinipes grasshoppers. Total shrub biomass significantly increased as carbon dioxide levels increased, as did the weight and area of individual leaves. Plants grown from seed collected in a single population exhibited a 3–5 fold variation in the concentration of leaf volatile mono- and sesquiterpenes, guaianolide sesquiterpene lactones, coumarins and flavones within each CO₂ treatment. The concentration of leaf allelochemicals did not differ significantly among CO₂ treatments for these seed-propagated plants. Further, when genotypic variation was controlled by vegetative propagation, allelochemical concentrations also did not differ among carbon dioxide treatments. On the other hand, overall leaf nitrogen concentration declined significantly with elevated CO₂. Carbon accumulation was seen to dilute leaf nitrogen as the balance of leaf carbon versus nitrogen progressively increased as CO₂ growth concentration increased. Grasshopper feeding was highest on sagebrush leaves grown under 270 and 650 μl l⁻¹ CO₂, but varied widely within treatments. Leaf nitrogen concentration was an important positive factor in grasshopper relative growth but had no overall effect on consumption. Potential compensatory consumption by these generalist grasshoppers was apparently limited by the sagebrush allelochemicals. Insects with a greater ability to feed on chemically defended host plants under carbon dioxide enrichment may ultimately consume leaves with a lower nitrogen concentration but the same concentration of allelochemicals. Compensatory feeding may potentially increase the amount of dietary allelochemicals ingested for each unit of nitrogen consumed.     

Effects of different macrophyte growth forms on sediment and P resuspension in a shallow lake

The effects of floating-leaved, submerged and emergent macrophytes on sediment resuspension and internal phosphorus loading were studied in the shallow Kirkkojärvi basin by placing sedimentation traps among different plant beds and adjacent open water and by sediment and water samples. All the three life forms considerably reduced sediment resuspension compared with non-vegetated areas. Both among submerged (Ceratophyllum demersum, Potamogeton obtusifolius, Ranunculus circinatus) and emergent (Typha angustifolia) plants, resuspension rate was on average 43% of that in the adjacent open water, while within floating-leaved plants (Nuphar lutea) the corresponding value was 87%. The effects of submerged and emergent vegetation increased in the course of the growing season together with increasing plant density. Among floating-leaved vegetation, such seasonal trend in resuspension effects was not observed. Compared with the non-vegetated area, floating-leaved, submerged and emergent plants reduced internal phosphorus loading on average by 21, 12 and 26 mg m⁻² d⁻¹, respectively. The effects of floating-leaved plants on resuspension-mediated internal phosphosrus loading were thus comparable to the effects of the other two life forms.     

Response of an insect herbivore to host plants grown in carbon dioxide enriched atmospheres

Rising atmospheric carbon dioxide concentration is expected to increase plant productivity, but little evidence is available regarding effects on insect feeding or growth. Larvae of the soybean looper, a noctuid moth, were fed leaves of soybean plants grown under three carbon dioxide regimes (350, 500 and 650 l·l^-1). Larvae fed at increasingly higher rates on plants from elevated carbon dioxide atmospheres: 30% greater on leaves from the 650 l·l^-1 treatment than on leaves from the 350 l·l^-1 treatment. When variation in larval feeding was related to the leaf content of nitrogen and water, there was no significant remaining effect of carbon dioxide treatment. The principal effect on herbivores of increasing the carbon supply of leaves appeared to be reduction of leaf nutrient concentration. This study suggests that feeding by herbivores on the leaves of C₃ plants may increase as the level of atmospheric carbon dioxide rises.     

Acquisition and assimilation of gaseous ammonia as revealed by intracellular pH changes in leaves of higher plants

Atmospheric ammonia (NH₃) from various anthropogenic sources has become a serious problem for natural vegetation. Ammonia not only causes changes in plant nitrogen metabolism, but also affects the acid-base balance of plants. Using the pH-sensitive fluorescent dyes pyranine and esculin, cytosolic and vacuolar pH changes were measured in leaves of C₃ and C₄ plants exposed for brief periods to concentrations of NH₃ in air ranging from 1.33 to 8.29 mol NH₃ · mol^-1 gas (0.94–5.86 mg · m^-3). After a lag phase, uptake of NH₃ from air at a rate of 200 nmol NH₃ · m ^{- 2} leaf area · s^{- 1} into leaves of Zea mays L. increased pyranine fluorescence indicating cytosolic alkalinisation. The increase was much larger in the dark than in the light. In illuminated leaves of the C₃ plant Pelargonium zonale L. and the C₄ plants Z. mays and Amaranthus caudatus L., NH₃-dependent cytosolic alkalinisation was particularly pronounced when CO₂ was supplied at very low levels (16 or 20 mol CO₂ · mol^{- 1} gas, containing 210 mmol O₂ · mol^{- 1} gas). An increase in esculin fluorescence, which was smaller than that of pyranine, was indicative of trapping of some of the NH₃ in the vacuoles of leaves of Spinacia oleracea L. and Z. mays. Photosynthesis and transpiration remained unchanged during exposure of illuminated leaves to NH₃, yielding an influx of 200 nmol NH₃ · m^-2 leaf area · s^-1 for up to 30 min, the longest exposure time used. Both CO₂ and O₂ influenced the extent of cytosolic alkalinisation. At 500 mol CO₂ · mol^-1 gas the cytosolic alkalinisation was suppressed more than at 16 or 20 mol CO₂ · mol^-1 gas. The suppressing effect of CO₂ on the NH₃induced alkalinisation was larger in illuminated leaves of the C₄ plants Z. mays and A. caudatus than in leaves of the C₃ plant P. zonale. A reduction of the O₂ concentration from 210 to 10 mmol O₂ · mol ^-1 gas, which inhibits photorespiration, increased the NH₃induced cytosolic alkalinisation in C₃ plants. Suppression by CO₂ or O₂ of the alkaline pH shift caused by the dissolution and protonation of NH₃ in queous leaf compartments, and possibly by the production of organic compounds synthesised from atmospheric NH₃, indicates that NH₃ which enters leaves is rapidly assimilated if photosynthesis or photorespiration provide nitrogen acceptor molecules.This work was supported by the Biotechnology and Biological Sciences Research Council and the Deutsche Forschungsgemein-schaft within the framework of the research of Sonderforschun-gsbreich 251 of the University of Würzburg. We are grateful to Dr. B. Wollenweber (The Royal Veterinary and Agricultural University, Denmark) for discussions.     

Acclimation to low light by C4 maize: implications for bundle sheath leakiness

C4 plants have a biochemical carbon concentrating mechanism (CCM) that increases CO₂ concentration around ribulose bisphosphate carboxylase oxygenase (Rubisco) in the bundle sheath (BS). Under limiting light, the activity of the CCM generally decreases, causing an increase in leakiness, (Φ), the ratio of CO₂ retrodiffusing from the BS relative to C4 carboxylation processes. Maize plants were grown under high and low light regimes (respectively HL, 600 versus LL, 100 μE m^?2 s^?1). Short‐term acclimation of Φ was compared from isotopic discrimination (Δ), gas exchange and photochemistry. Direct measurement of respiration in the light, and ATP production rate (J_ATP), allowed us use a novel approach to derive Φ, compared with the conventional fitting of measured and predicted Δ. HL grown plants responded to decreasing light intensities with the well‐documented increase in Φ. Conversely, LL plants showed a constant Φ, which has not been observed previously. We explain the pattern by two contrasting acclimation strategies: HL plants maintained a high CCM activity at LL, resulting in high CO₂ overcycling and increased Φ; LL plants acclimated by down‐regulating the CCM, effectively optimizing scarce ATP supply. This surprising plasticity may limit the impact of Φ‐dependent carbon losses in leaves becoming shaded within developing canopies.     

Effects of Irradiance-Sulphur Interactions on Enzymes of Carbon,Nitrogen, and Sulphur Metabolism in Maize Plants

The effect of sulphur deprivation and irradiance (180 and 750 µmol m^–2 s^–1) on plant growth and enzyme activities of carbon, nitrogen, and sulphur metabolism were studied in maize (Zea mays L. Pioneer cv. Latina) plants over a 15-d-period of growth. Increase in irradiance resulted in an enhancement of several enzyme activities and generally accelerated the development of S deficiency. ATP sulphurylase (ATPs; EC 2.7.7.4) and o-acetylserine sulphydrylase (OASs; EC 4.2.99.8) showed a particular and different pattern as both enzymes exhibited maximum activity after 10 d from the beginning of deprivation period. Hence in maize leaves the enzymes of C, N, and S metabolism were differently regulated during the leaf development by irradiance and sulphur starvation.     

Procedure for determination of elemental sulphur bio-oxidation rate

A procedure is described for measuring the rate of biooxidation of elemental sulphur in nutrient solutions. Results of preliminary measurements of sulphur bio-oxidation rate in a dynamic system are presented. The rate of sulphur bio-oxidation has been determined at the level of 0.02–0.05 g of sulphur per m² of sulphur per h.List of Symbols C g/dm³ concentration of sulphate ions - C ₂ g/dm³ concentration of sulphate ions in withdrawn solution - C g/dm³ C difference between solution outlet and inlet to sulphur bed - F m² sulphur surface exposed to bacteria action - m g mass of elemental sulphur - V dm³ volume of solution - V ₀ dm³/h volume of fresh solution supplied to the set - V ₁ dm³/h circulating solution flow rate - V ₂ dm³/h volume of solution withdrawn - h time Abbreviations RBES rate of bio-oxidation of elemental sulphur     

二氧化硫污染对植物体酶活性的影响

通过用６种浓度的ＳＯ２气体对６种植物进行熏气处理,然后测定植物叶片内的过氧化物酶,抗坏血酸氧化酶及多酚氧化酶的活性,为环境监测提供理论依据。通过试验,发现ＳＯ２气体对植物叶内不同酶的活性的影响有别。６种植物受不同浓度的ＳＯ２污染,叶内的过氧化物酶活性随ＳＯ２浓度增加而增加,促进叶片的衰老;而对其他的２种酶,则６种植物表现不同     

Photosynthetic activity of poikilochlorophyllous desiccation tolerant plant <Emphasis Type="Italic">Reaumuria soongorica</Emphasis> during dehydration and re-hydration

Diurnal patterns of gas exchange and chlorophyll (Chl) fluorescence parameters of photosystem 2 (PS2) as well as Chl content were analyzed in Reaumuria soongorica (Pall.) Maxim., a perennial semi-shrub during dehydration and rehydration. The net photosynthetic rate (P _N), maximum photochemical efficiency of PS2 (variable to maximum fluorescence ratio, F_v/F_m), quantum efficiency of non-cyclic electron transport of PS2, and Chl content decreased, but non-photochemical quenching of fluorescence and carotenoid content increased in stems with the increasing of drought stress. 6 d after re-hydration, new leaves budded from stems. In the re-watered plants, the chloroplast function was restored and Chl a fluorescence returned to a similar level as in the control plants. This improved hydraulic adjustment in plant triggered a positive effect on ion flow in the tissues and increased shoot electrical admittance. Thus R. soongorica plants are able to sustain drought stress through leaf abscission and keep part of Chl content in stems.     

The Transport of Oleanolic Acid in Calendula officinalis

The transport of oleanolic acid synthesized from ¹⁴CO₂ in the individual leaves of Calendula officinalis plants was investigated. It was proved that the rate and the direction of the transport depend on the stage of vegetation and the position of the leaf on the plant. The greatest transport rates of oleanolic acid were observed from the upper level leaves towards the inflorescence during the flowering period. The overblowing period was characterized by a slower migration of oleanolic acid from the leaf supplied with radioactive precursor. The migration was directed mainly to the root especially from the low level leaves. Using CH₃¹⁴COONa as precursor, it was established that the root is unable to synthesize oleanolic acid.     

ATP and NADPH as the driving force of carbon reduction in leaves in relation to thylakoid energization by light

Carbon assimilation of spinach (Spinacia oleracea L.) leaves was measured in the presence of 2000l· l^–1CO₂ and 2% O₂ in the gas phase to suppress photorespiratory reactions and to reduce stomatal diffusion resistance. Simultaneously, membrane parameters such as modulated chlorophyll fluorescence, oxidation of P₇₀₀ in the reaction centre of photosystem I, and apparent changes in absorbance at 535 nm were recorded. After light-regulated enzymes were activated at a high irradiance, illumination was changed. About 3 min later (to maintain the previous activation state of enzymes), leaves were shock-frozen and freeze-dried. Chloroplasts were isolated nonaqueously and analysed for ATP, ADP, inorganic phosphate, NADPH and NADP. Observations made under the chosen conditions differed in some important aspects from those commonly observed when leaves are illuminated in air. (i) Not only assimilation, but also the phosphorylation potential [ATP]/([ADP]·[Pi]) increased hyperbolically with irradiance towards saturation. In contrast, the ratio of NADPH to NADP did not change much as irradiances increased from low to high photon flux densities. When ATP, the phosphorylation potential and the assimilatory force, F_A (the product of phosphorylation potential and NADPH/NADP ratio), were plotted against assimilation, ATP increased relatively less than assimilation, whereas the phosphorylation potential increased somewhat more steeply than assimilation did. A linear relationship existed between assimilation and F_A at lower irradiances. The assimilatory force F_A increased more than assimilation did when irradiances were very high. Differences from previous observations, where F_A was under some conditions higher at low than at high rates of carbon assimilation, are explained by differences in flux resistances caused not only by stomatal diffusion resistance but also by differences in the activity of light-regulated enzymes, (ii) The relationship between P₇₀₀ oxidation and a fast absorption change with a maximum close to 520 nm on one hand and carbon assimilation on the other hand was largely linear under the specific conditions of the experiments. A similar linear relationship existed also between the quantum efficiency of electron flow through photosystem II and the quantum efficiency of photosystem I electron transport. (iii) Whereas the increase in non-photochemical fluorescence quenching, q_N, was similar to the increase in assimilation, the relationship between light scattering and assimilation was distinctly sigmoidal. Light scattering appeared to be a better indicator of control of photosystem II activity under excessive irradiation than q_N. (iv) The results are discussed in relation to the relative significance of chloroplast levels of ATP and NADPH and of the assimilatory force F_A in driving carbon assimilation. From the observations, the proton/electron (H⁺/e^–) ratio of linear electron transport is suggested to be 3 and the H⁺/ATP ratio to be 4 in leaves. An H⁺/e^– ratio of 3 implies the existence of an obligatory Q-cycle in leaves.Abbreviations F_A assimilatory force - F_o fluorescence after long dark adaptation - F_m maximum fluorescence level - F_s steady-state fluorescence - PGA 3-phosphoglycerate - PFD photon flux density - P₇₀₀ (P₇₀₀+) electron-donor pigment in the reaction center of PSI (its oxidized form) - Q_A primary quinone acceptor of PSII - q_P photochemical quenching - q_N non-photochemical quenching - _PSII relative quantum efficiency of energy conversation at the level of photosystem II - _PSI relative quantum efficiency of photosystem II This research was supported by the Sonderforschungsbereich 251 of the University of Würzburg and the Stiftung Volkswagenwerk. U.G. is a member of the Graduate College of the Julius-von-Sachs Institut für Biowissenschaften, University of Würzburg, being on leave from Tartu University, Tartu, Estonia. The authors are grateful to Prof. A. Laisk, Chair of Plant Physiology, Tartu University, for stimulating discussions.     

Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light

Diurnal changes in photosynthetic gas exchange and chlorophyll fluorescence were measured under full sunlight to reveal diffusional and non‐diffusional limitations to diurnal assimilation in leaves of Arisaema heterophyllum Blume plants grown either in a riparian forest understorey (shade leaves) or in an adjacent deforested open site (sun leaves). Midday depressions of assimilation rate (A) and leaf conductance of water vapour were remarkably deeper in shade leaves than in sun leaves. To evaluate the diffusional (i.e. stomatal and leaf internal) limitation to assimilation, we used an index [1–A/A₃₅₀], in which A₃₅₀ is A at a chloroplast CO₂ concentration of 350 μ mol mol ^{? 1}. A₃₅₀ was estimated from the electron transport rate (J_T), determined fluorometrically, and the specificity factor of Rubisco (S), determined by gas exchange techniques. In sun leaves under saturating light, the index obtained after the ‘peak’ of diurnal assimilation was 70% greater than that obtained before the ‘peak’, but in shade leaves, it was only 20% greater. The photochemical efficiency of photosystem II ( Δ F/F_m ′ ) and thus J_T was considerably lower in shade leaves than in sun leaves, especially after the ‘peak’. In shade leaves but not in sun leaves, A at a photosynthetically active photon flux density (PPFD) > 500 μ mol m ^{? 2} s ^{? 1} depended positively on J_T throughout the day. Electron flows used by the carboxylation and oxygenation (J_O) of RuBP were estimated from A and J_T. In sun leaves, the J_O/J_T ratio was significantly higher after the ‘peak’, but little difference was found in shade leaves. Photorespiratory CO₂ efflux in the absence of atmospheric CO₂ was about three times higher in sun leaves than in shade leaves. We attribute the midday depression of assimilation in sun leaves to the increased rate of photorespiration caused by stomatal closure, and that in shade leaves to severe photoinhibition. Thus, for sun leaves, increased capacities for photorespiration and non‐photochemical quenching are essential to avoid photoinhibitory damage and to tolerate high leaf temperatures and water stress under excess light. The increased Rubisco content in sun leaves, which has been recognized as raising photosynthetic assimilation capacity, also contributes to increase in the capacity for photorespiration.