Photosynthetic Electron Transport in Shoots of Sitka Spruce from Different Levels in a Forest Canopy

Shoots from the lower levels of a Sitka forest canopy [Picea sitchensis (Bong.) Carr.] have much lower photosynthetic capacities per unit leaf area than do those from the top of the canopy. Chloroplast fragments have been isolated from the needles at three levels in the canopy and Photosystem I and Photosystem II activities measured. Measurements were made at eight photon flux densities giving light response curves for photosystem activity in the three canopy levels. Expressed per unit leaf area, light-saturated activities of Photosystem I and II were lower at the bottom of the canopy than at the top suggesting that photosystem activity might be limiting photosynthesis in the shoots from the bottom of the canopy. Chlorophyll concentration per unit fresh weight did not vary amongst the three levels, but specific leaf area increased and dry weight fraction decreased with increasing depth in the canopy, leading to a large reduction with depth in chlorophyll content per unit leaf area.     

Effects of natural shade on soybean thylakoid membrane composition

The effect of natural shade on chloroplast thylakoid membrane activity and composition was examined for soybean (Glycine Max. cv. Young) grown under field conditions. Plots with high (10 plants m^–1 row) or low (1 plant m^–1 row) plant density were established. Expanding leaves were tagged at 50, 58 and 65 days after planting (DAP). At 92 DAP, tagged leaves were used as reference points to characterize canopy light environments and isolate thylakoid membranes. Light environments ranged from a photosynthetic photon flux density (PPFD) of 87% of full sun to a PPFD of 10% of full sun. The decline in PPFD was accompanied by an increase in the far-red/red (735 nm/645 nm) ratio from 0.9 to approximately six. The major effects of shade on chloroplast thylakoid membranes were a reduction in chloroplast coupling factor and a shift in light-harvesting capacity from Photosystem I to Photosystem II. Photosynthetic electron transport capacity was not affected by differences in PPFD, but was 20 to 30% higher in the 1 plant m^–1 row treatment. The plant density effect on electron transport was associated with differences in plastocyanin concentration, suggesting that plastocyanin is a limiting factor in soybean. Shade did not have a significant effect on the concentration of Photosystem II, Cyt b₆f, or Photosystem I complexes.Abbreviations CF₁ chloroplast coupling factor - DAP days after planting - DBMIB 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone - DCIP 2,6-dichlorophenolindophenol - FR/R far-red/red - PBS 10 mM sodium phosphate (pH 7.0), 150 mM NaCl - PPFD photosynthetic photon flux density - PS I Photosystem I - PS II Photosystem II - P700 reaction center of Photosystem I - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - TBS 20 mM Tris-HCl (pH 7.5), 500 mM NaCl - TTBS 20 mM Tris-HCl (pH 7.5), 500 mM NaCl, 0.05% (w/v) polyoxyethylenesorbitan monolaurate (Tween-20) The US Government right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged.The US Government right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged.     

Changes in composition and function of thylakoid membranes as a result of photosynthetic adaptation of chloroplasts from pea plants grown under different light conditions

The hypothesis that chloroplasts having different light-saturated rates of photosynthesis will have different proportions of the intrinsic thylakoid complexes engaged in light-harvesting and electron transport (Anderson, J.M. (1982) Mol. Cell. Biochem. 46, 161–172) has been tested. Peas were grown in light regimes which varied in light intensity, quality and time of irradiance, and ranged from sunlight through red to blue-enriched light of very low radiation. The electron-transport capacity at saturating light of Photosystem I and Photosystem II of chloroplasts isolated from light-adapted peas was 2-fold and 5–6-fold lower, respectively, in the lowest radiation compared to sunlight. There was a marked increase in the amount of total chlorophyll associated with the main chlorophyll $\text{a}\text{b-}\text{proteins}$ (LHCP¹, LHCP² and LHCP³) and a 2-fold decrease in the core reaction centre complex of Photosystem II (CP a) as the radiation decreased; the $\text{LHCP}{}^{\mathrm{1–3}}\text{CP}$ a ratio changed from 3.5 to 9.0. The amount of chlorophyll associated with Photosystem I varied from 34% in sunlight to 27% in the lowest radiation, but the antenna size of Photosystem I was not markedly different; there was a 2-fold decrease in the amount of cytochrome f on a chlorophyll basis, which partly accounted for the decreased electron-transport capacity of Photosystem I. Since the increases or decreases in the levels of each of the components correlated with decreasing radiation, it is clear that the light-adaptation of both light-harvesting and electron-transport components is indeed closely co-ordinated.     

Quantum Requirements of Photosynthetic Electron Transport In Sitka Spruce from Different Light Environments

Quantum requirements of photosynthetic electron transport have been measured in shoots of Picea sitchensis (Bong.) Carr. (Sitka spruce) from different levels in a forest canopy and in shoots from plants grown in contrasting light environments in controlled environment chambers. Neutral density filters were used to obtain very low photon flux densities. The light absorbed by the chloroplast suspensions was calculated from measurements of the transmittance of the suspensions. The shoots from the top of the forest canopy (“sun” shoots) had lower quantum requirements for photosystems I and II than the shoots from the bottom of the forest canopy (“shade” shoots). High light grown plants and “sun” shoots had higher rates of electron transport at light saturation than low light grown plants and “shade” shoots. Thus a higher potential for electron transport was found to exist in “sun” shoots than in “shade” shoots at both high and low photon flux densities.     

Light adaptation of cyclic electron transport through Photosystem I in the cyanobacterium Synechococcus sp. PCC 7942

Photosystem I-driven cyclic electron transport was measured in intact cells of Synechococcus sp PCC 7942 grown under different light intensities using photoacoustic and spectroscopic methods. The light-saturated capacity for PS I cyclic electron transport increased relative to chlorophyll concentration, PS I concentration, and linear electron transport capacity as growth light intensity was raised. In cells grown under moderate to high light intensity, PS I cyclic electron transport was nearly insensitive to methyl viologen, indicating that the cyclic electron supply to PS I derived almost exclusively from a thylakoid dehydrogenase. In cells grown under low light intensity, PS I cyclic electron transport was partially inhibited by methyl viologen, indicating that part of the cyclic electron supply to PS I derived directly from ferredoxin. It is proposed that the increased PSI cyclic electron transport observed in cells grown under high light intensity is a response to chronic photoinhibition.Abbreviations DBMIB 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone - DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea - ES energy storage - MV methyl viologen - PA_m photoacoustic thermal signal with strong non-modulated background light added - PA_s photoacoustic thermal signal without background light added CIW/DPB Publication No. 1205.     

Effect of growth temperature on the composition of the photosynthetic apparatus in a thermophilic cyanobacterium (Synechococcus sp.)

Abstract A thermophilic strain of Synechococcus sp. was grown under light-limited conditions at its optimum temperature of 58°C and at 38°C in order to investigate the effect of growth temperature on the composition of the photosynthetic apparatus. Cells grown at 38°C had a ratio of Photosystem I to Photosystem II of 2.2, close to that known to occur in unstressed mesophilic strains of Synechococcus . Growth at the higher temperature led to a doubling of the number of Photosystem I reaction centres per cell whilst Photosystem II remained essentially unchanged, causing the Photosystem I/Photosystem II ratio to rise to 4.1. These changes were correlated with an increase in the number of concentric layers of thylakoid membranes from four to nine. It is suggested that these changes result from a higher relative demand for energy (ATP) during growth at elevated temperatures.     

Effect of growth irradiance on plastocyanin levels in barley

Plastocyanin levels in barley (Hordeum vulgare cv Boone) were found to be dependent on growth irradiance. An immunochemical assay was developed and used to measure the plastocyanin content of isolated thylakoid membranes. Barley grown under 600 mole photons m^–2s^–1 contained two- to four-fold greater quantities of plastocyanin per unit chlorophyll compared with plants grown under 60 mole photons m^–2s^–1. The plastocyanin/Photosystem I ratio was found to be 2 to 3 under high irradiance compared with 0.5 to 1.5 under low irradiance. The reduced plastocyanin pool size in low light plants contributed to a two-fold reduction in photosynthetic electron transport activity. Plastocyanin levels increased upon transfer of low light plants to high irradiance conditions. In contrast, plastocyanin levels were not affected in plants transferred from high to low irradiance, suggesting that plastocyanin is not involved in the acclimation of photosynthesis to shade.Abbreviations: BSA bovine serum albumin - chl chlorophyll - cyt cytochrome - DCIP 2,6-dichlorophenolindophenol - PS I Photosystem I - PS II Photosystem II - P700 reaction center of Photosystem I - TBS 20 mM Tris-HCl pH 7.5, 500 mM NaCl - TTBS 20 mM Tris-HCl pH 7.5, 500 mM NaCl, 0.5% (w/v) polyoxyethylenesorbitan monolaurate (Tween-20)     

Increase in Nitrate Reductase Activity in Bean Leaves by Light Involves a Regulator Protein

Nitrate reductase activity, assayed either in vivo or in vitro was considerably higher in bean (Phaseolus vulgaris L.) leaves from 7-day-old light grown seedlings than those from dark grown, both in the absence as well as presence of nitrate. Cytochrome c reductase activity was however similar in both regimes, while peroxidase was lower in light than in dark. The light stimulated increase in nitrate reductase activity in leaf segments from dark grown seedlings was inhibited by cycloheximide, DNP, chloramphenicol, and sodium tungstate and was unaffected by lincomycin and DCMU. Under similar conditions, the increase in total chlorophyll was inhibited completely by cycloheximide and DNP, partially by chloramphenicol and lincomycin, and was unaffected by tungstate and DCMU. A supply of 1~5 mm reduced glutathione increased enzyme activity in the dark and also to some extent in light. The substrate induction of enzyme activity started after a lag of one hr in light or dark and continued for either 5 hr in the dark or 8 hr in light. Two proteinaceous inhibitors (Factors I and II) of nitrate reductase were isolated by ammonium sulfate precipitation and Sephadex gel filtration. The amount of Factor I was higher in the dark than in light. The amount and activity of Factor II was however, almost equal in light and dark. The inhibition of enzyme activity by these inhibitors increased with their concentration. It is proposed that light increases nitrate reductase activity by decreasing the amount of a nitrate reductase inhibitor.     

Relation of light and nitrogen source to growth, nitrate reductase and glutamine synthetase activity of jack pine seedlings

Two-month-old jack pine ( Pinus banksiana Lamb.) seedlings were placed in a greenhouse where both nitrogen source and light level were varied. After 4 months, whole seedling biomass, leaf biomass and relative growth rate were greatest in seedlings grown with NH⁺₄/NO/NO⁻₃-N and full light (FL) and least in seedlings grown with NO ⁻₃-N and low light (LL). NO ⁻₃-seedlings grown under full light and NH⁺₄/NO⁻₃-seedlings grown under low light were approximately equal. This indicates that the extra carbon costs of assimilating only NO⁻₃-N were similar to the reduction of carbon fixation resulting from a 50% decrease in photon flux density. Percentage and total nitrogen content of needles were greater in seedlings grown under low light independent of nitrogen fertilization. Percentage and total nitrogen content of roots were higher under low light and lower when fertilized with NO⁻₃.
Nitrate reductase (NR) activity was higher in roots than in needles, while glutamine synthetase (GS) activity was higher in needles than in roots. Low light resulted in decreased NR activity (mg N)⁻¹ in needles, but not in roots. However, no nitrate was detected in the needles in any treatment. GS activity, on the other hand, was greater under low light in both needles and roots. GS activity in needles is most likely involved with the reassimilation rather than the initial assimilation of ammonium. Some implications of these shifts in enzymatic activity for ecological phenomena in forests are discussed.     

Prolonged high light treatment of plant cells results in changes of the amount,the localization and the electrophoretic behavior of several thylakoid membrane proteins

The effect of a 30 h high light treatment on the amount and the localization of thylakoid proteins was analysed in low light grown photoautotrophic cells of Marchantia polymorpha and Chenopodium rubrum. High light treatment resulted in a net loss of D1 protein which was accompanied by comparable losses of other proteins of the PS II core (reaction center with inner antenna). LHC II proteins were not reduced correspondingly, indicating that these complexes are less affected by prolonged high light. High light influenced the distribution of PS II components between the grana and the stroma region of the thylakoid membrane, probably by translocation of the respective PS II proteins. Additionally, modifications of several thylakoid proteins were detected in high light treated cells of C. rubrum. These effects are discussed in relation to photoinhibitory damage and repair processes.Abbreviations BCA bioinchonic acid - chl chlorophyll - CF₁ coupling factor - CYC cycloheximide - GT grana thylakoids - HL high light - LL low light - PAGE polyacrylamide gel electrophoresis - PFD photon flux density - PS I Photosystem I - PS II Photosystem II - RC reaction center - SDS sodium dodecylsulfate - ST stroma thylakoids - Thyl unfractionated thylakoids     

Photosystem 1 preparations from dark-and light-grown cells of the cyanobacterium,Chlorogloea fritschii

Photosystem 1 (PS1) enriched preparations have been extracted from the cyanobacterium Chlorogloea fritschii grown either in darkness or in the light. Absorption spectra show that the main chlorophyll peak has shifted from 678 nm in PS1 from light grown cells to 675 nm in PS1 from dark grown cells. Fluorescence spectra show a similar blue shift in wavelength maximum from 690 nm to 678 nm and the fluorescence intensity is higher in PS1 from dark grown cells. Allophycocyanin is present in PS1 from light grown cells, but absent from preparations from C. fritschii grown in the dark. P700: chlorophyll a ratios of the preparations from light and dark grown cells are 1:35 and 1:80 respectively, all P700 being photoactive. The results are interpreted to suggest that allophycocyanin is not attached to PS1 in dark grown C. fritschii, neither is all chlorophyll arranged in such a way as to ensure efficient energy transfer to P700.     

The far red induced slow component of delayed light from chloroplasts is emitted from Photosystem II

In spinach chloroplasts illuminated with far red light, the relative intensity maximum during the decay of delayed light is emitted at 680–690 nm. This finding supports previous models predicting emission from Photosystem II, and contradicts earlier attributions to Photosystem I.Due to self absorption, the emission spectrum of the relative maximum is shifted to longer wavelengths and displays apparent Photosystem I characteristics in chloroplast samples of higher concentration or in leaves. This may have caused earlier investigators to ascribe the emission to Photosystem I.A differences between the spectral width of the emission spectra of delayed fluorescence and the relative maximum indicates that these two phenomena represent emission from different sub-populations of Photosystem II centers.Abbreviations PS I Photosystem I - PS II Photosystem II - DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea     

The Spectral and Photochemical Properties of Subchloroplast Membrane Particles from Pinus silvestris

Isolated chloroplasts from Pinus silvestris have been fractionated by a combination of digitonin and Yeda-press treatment. Different subchloroplast particles have been isolated by differential centrifugation. The spectral and photochemical properties of the different fractions have been analysed. Photosystem I was enriched in the light particles (high Chl a/Chl b ratios, low Chl/P700 ratios, high F735/F685 ratios, high photosystem I activity and enrichment of long-wavelength absorbing Chl a). The heavy fractions were enriched in photosystem II (low Chl a/Chl b ratios, high Chl/P700 ratios, low F735/F685 ratios and enrichment of short-wavelength absorbing Chl a). The molar ratio Chl/P700 was about 1.5 times greater for Pinus than for Spinacia, and Pinus had relatively less long-wavelength absorbing Chl a compared with Spinacia.     

Effect of the respiratory activity on photoinhibition of the cyanobacterium Synechocystis sp.

The influence of respiratory activity on photosynthesis in Synechocystis cells that had been exposed to high light intensity was studied using distinct conditions of nitrogen supply. The photoinhibitory rate of N-sufficient cells was not influenced by the presence of different nitrogen sources. In contrast, when N-starved cells were resupplied with ammonium, they were protected from photoinhibition. Although N-starved cells presented a higher rate of dark O₂ uptake than N-sufficient ones, the photoinhibitory rate increased in both cases after addition of sodium azide or sodium azide plus salicylhydroxamic acid in the photoinhibitory treatment. In the absence of the D₁ protein repair mechanism, photodamage to Photosystem II was faster in N-sufficient cells than in N-starved ones. Mitigation of photodamage disappeared when the respiratory activity of N-starved cells was partially suppressed by the addition of sodium azide or sodium azide and salicylhydroxamic acid. Our results suggest that electron flow through cyanobacterial terminal oxidases can assist Photosystem I in removing electrons from the reduced plastoquinone pool, thus contributing to both reopening of Photosystem II reaction centers and avoiding photogeneration of reactive oxygen species under photoinhibitory conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

Determination of the antenna heterogeneity of Photosystem II by direct simultaneous fitting of several fluorescence rise curves measured with DCMU at different light intensities

Several methods for determination of the antenna heterogeneity of Photosystem II from fluorescence rise curves measured with DCMU have been developed so far. Using these methods, two, three or four types of Photosystem II with respect to the antenna heterogeneity were determined. However, the accuracy of some of these methods is under debate. Here, we present a new method for the determination of the antenna heterogeneity of Photosystem II. The method is based on direct simultaneous fitting of several fluorescence rise curves measured with DCMU at different intensities of light excitation. As several curves measured under different light conditions are fitted simultaneously by the same model, reliability and accuracy in determination of model parameters increase. Our method was applied to two plant materials with different structure of the thylakoid membrane: wheat leaves and cells of green alga Chlamydomonas reinhardtii. This revised version was published online in June 2006 with corrections to the Cover Date.     

The influence of light on the growth of watercress (<Emphasis Type="Italic">Nasturtium officinale</Emphasis> R. Br.)

In stream ecosystems, the growth of aquatic primary producers is affected by spatial and temporal variations in the riparian canopy, which can influence the availability of light resources. Aquatic plants can acclimate to low light environments by employing a suite of morphological or physiological mechanisms to increase light capture or photosynthetic efficiency. Some species may also use alternate types of propagules to colonize environments with heterogeneous light environments. In a greenhouse experiment we examined the morphological and physiological response of watercress (Nasturtium officinale R. Br.) to a gradient of increasing light levels, which ranged from 7% ambient light to full sunlight. We also determined if watercress seedlings and vegetative fragments differed in their growth response to increasing light levels. Total biomass and root biomass of seedlings and vegetative fragments decreased with decreasing light levels. The difference in plant biomass across treatments was due to morphological changes in total canopy area and leaf area, both of which increased with decreasing light levels. Seedlings and vegetative fragments did not differ in their response to light availability, but vegetative fragments had higher final biomass as a result of higher initial biomass. Physiological acclimation to low light levels appears to be of secondary importance for watercress as the concentrations of total chlorophyll, chlorophyll a, chlorophyll b, and chlorophyll a:b did not differ among light levels or between seedlings and vegetative fragments. Seedlings and vegetative fragments grown under high light levels had a greater percentage of carbon and a lower percentage of nitrogen than plants grown under low light conditions. The results of this study indicate that watercress displays considerable morphological plasticity and acclimates to low light conditions primarily by increasing leaf area and canopy surface area. There is no evidence that the type of watercress propagule (seedling vs. vegetative fragment) imparts any growth advantage in low light environments and watercress grown from either type of propagule showed no differences in their morphological or physiological responses to varying light regimes. Handling editor: S. M. Thomaz     

The influence of high levels of brief or prolonged supplementary far-red illumination during growth on the photosynthetic characteristics, composition and morphology of Pisum sativum chloroplasts

Abstract. Peas were grown in controlled environments (12h white fluorescent light. ∼47 μmol photons m^-2 s 1/12 dark, 25 °C), using (1) 15-min far-red illumination at the end of each photoperiod (brief FR) to simulate the increase in the far-red/red ratio near the end of the day, and (2) high levels of supplementary far-red light (red:far-red ratio=0.04) during the entire photoperiod (long-term FR) to simulate extreme shade conditions under a plant canopy. Brief FR illumination led to marked morphological effects attributable to phytochrome regulation, namely, an increase in internodal length, but a decrease in leaflet area, chloroplast size and chlorophyll content per chloroplast compared with the control. Significantly, brief FR illumination had little or no effect on the amounts of the major chloroplast components (ribulose 1.5-biphosphate carboxylase, adenosine triphosphate synthase, cytochrome b/f complex and Photosystem II) relative to chlorophyll or Photosystem I, and the leaf photosynthetic capacities per unit chlorophyll were similar. In contrast, supplementing high levels of far-red light during the entire photoperiod not only led to the phytochrome effects above, but there was also a marked increase in leaf photosynthetic capacity per unit chlorophyll. due to increased amounts of the major chloroplast components relative to chlorophyll or Photosystem I. We hypothesize that supplementary far-red light, absorbed by Photosystem I, induced an increase in the major chloroplast components by a photosynthetic feedback mechanism. In fully greened leaves, we propose that the two photosystems themselves, rather than phytochrome, may be the predominent sensors of light quantity in triggering modulations of the stoichiometries of chloroplast components, which in turn lead to varying photosynthetic capacities.     

Influence of light intensity during growth on photosynthesis and activity of several key photosynthetic enzymes in a C4 plant (Zea mays)

Maize ( Zea mays L. Hybrid Sweet Corn, Royal Crest), a C₄ plant, was grown under different light regimes, after which the rate of photosynthesis and activities of several photosynthetic enzymes (per unit leaf chlorophyll) were measured at different light intensities. Plants were grown outdoors under direct sunlight or 23% of direct sunlight, and in growth chambers at photosynthetic photon flux densities of about 20% and 8% of direct sunlight. The plants grown under direct sunlight had a higher light compensation point than plants grown under lower light. At a light intensity about 25% of direct sunlight, plants from all growth regimes had a similar rate of photosynthesis. Under saturating levels of light the plants grown under direct sunlight had a substantially higher rate of photosynthesis than plants grown under the lower light regimes. The higher photosynthetic capacity in the plants grown under direct sunlight was accompanied by an increased activity of several photosynthetic enzymes and in the amount of the soluble protein in the leaf. Among five photosynthetic enzymes examined, RuBP carboxylase (EC 4.1.1.39) and pyruvate, Pi dikinase (EC 2.7.9.1) were generally just sufficient to account for rates of photosynthesis under saturating light; thus, these may be rate limiting enzymes in C₄ photosynthesis. Pyruvate, Pi dikinase and NADP-malate dehydrogenase (EC 1.1.1.82) were the only enzymes examined which were light activated and increased in activity with increasing light intensity. In the low light grown plants the activity of pyruvate, Pi dikinase closely paralleled the photosynthetic rate measured under different light levels. With the plants grown under direct sunlight, as light intensity was increased the activation of pyruvate, Pi dikinase and NADP⁺-malate dehydrogenase proceeded more rapidly than photosynthesis.     

Stimulation of electron transport from Photosystem II to Photosystem I in spinach chloroplasts

Electron transport from Photosystem II to Photosystem I of spinach chloroplasts can be stimulated by bicarbonate and various carbonyl or carboxyl compounds. Monovalent or divalent cations, which have hitherto been implicated in the energy distribution between the two photosystems, i.e., spillover phenomena at low light intensities, show a similar effect under high light conditions employed in this study. A mechanism for this stimulation of forward electron transport from Photosystem II to Photosystem I could involve inhibition of two types of Photosystem II partial reactions, which may involve cycling of electrons around Photosystem II. One of these is the DCMU-insensitive silicomolybdate reduction, and the other is ferricyanide reduction by Photosystem II at pH 8 in the presence of dibromothymoquinone. Greater stimulation of forward electron transport reactions is observed when both types of Photosystem II cyclic reactions are inhibited by bicarbonate, carbonyl and carboxyl-type compounds, or by certain mono- or divalent cations.Abbreviations used: DCMU, 3-(3,4-dichlorophenyl)-1, 1-dimethylurea; DCIP, 2,6-dichloroindophenol; DBMIB, 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone; FeCN, potassium ferricyanide; MV, methylviologen; PS I, photosystem I; PS II, photosystem II; SM, silicomolybdic acid.