Action potential in a plant cell lowers the light requirement for non-photochemical energy-dependent quenching of chlorophyll fluorescence

This study deals with effects of membrane excitation on photosynthesis and cell protection against excessive light, manifested in non-photochemical quenching (NPQ). In Chara corallina cells, NPQ and pericellular pH displayed coordinated spatial patterns along the length of the cell. The NPQ values were lower in H⁺-extruding cell regions (external pH ∼ 6.5) than in high pH regions (pH ∼ 9.5). Generation of an action potential by applying a pulse of electric current caused NPQ to increase within 30-60 s. This effect, manifested as a long-lived drop of maximum chlorophyll fluorescence (F_m′), occurred at lower photosynthetic flux densities (PFD) in the alkaline as compared to acidic cell regions. The light response curve of NPQ shifted, after generation of an action potential, towards lower PFD. The release of NPQ by nigericin and the rapid reversal of action potential-triggered NPQ in darkness indicate its relation to thylakoid ΔpH. Generation of an action potential shortly after darkening converted the chloroplasts into a latent state with the F_m identical to that of unexcited cells. This state transformed to the quenched state after turning on weak light that was insufficient for NPQ prior to membrane excitation of the cells. The ionophore, A23187, shifted NPQ plots similarly to the action potential effect, consistent with a likely role of a rise in the cytosolic Ca²⁺ level in the action potential-induced quenching. The results suggest that a rapid electric signal, across the plasma membrane, might exert long-lived effects on photosynthesis and chlorophyll fluorescence through ion flux-mediated pathways.     

High-temperature damage and acclimation of the photosynthetic apparatus

The influence of mono- (K⁺) and divalent (Mg²⁺) cations and protons (pH) on the temperature sensitivity of thylakoid membranes was investigated in three groups of young bean plants (control, heat-acclimated and non-acclimated). Thylakoid-membrane function was monitored by second and millisecond delayed fluorescence and 9-aminoacridine fluorescence quenching. It was established that metal ions at investigated concentrations decreased the thermostability of the photosynthetic parameters — an increase of MgSO₄ concentration from 0.1 to 20 mM decreased the temperature of their half-inactivation (T50) by 13°C. At the same time the pH dependence of the thermal stability of these parameters showed a maximum at pH 5.5–6.5. The half-inactivation temperatures of those photosynthetic parameters connected with the ability of the thylakoid membrane to form light-induced proton gradients increased by 6–7°C in the heat-acclimated plants compared with the control. It was assumed that the temperature inactivation of photosynthetic electron transfer and the energization of the thylakoid membrane was determined both by the thermoinduced dissociation of the light-harvesting chlorophyll a/b protein complex from PSII, leading to destruction of the excitation energy transfer to the reaction centres, and by the thermal denaturation of the membrane-protein components. The rate of these processes was probably controlled by the size of the negative surface charge and the viscosity of the thylakoid membrane.Abbreviations 9-AA 9-aminoacridine - DF delayed fluorescence - LHCP light-harvesting chlorophyll a/b protein complex - PSI (II) photosystem I (II) - T50 temperature of 50% inhibition of photosynthetic parameter - Tricine N-[2-hydroxy-1, 1-bis(hydroxymethyl)ethyl] glycine     

Action potential in a plant cell lowers the light requirement for non-photochemical energy-dependent quenching of chlorophyll fluorescence

This study deals with effects of membrane excitation on photosynthesis and cell protection against excessive light, manifested in non-photochemical quenching (NPQ). In Chara corallina cells, NPQ and pericellular pH displayed coordinated spatial patterns along the length of the cell. The NPQ values were lower in H(+)-extruding cell regions (external pH approximately 6.5) than in high pH regions (pH approximately 9.5). Generation of an action potential by applying a pulse of electric current caused NPQ to increase within 30-60 s. This effect, manifested as a long-lived drop of maximum chlorophyll fluorescence (F(m)'), occurred at lower photosynthetic flux densities (PFD) in the alkaline as compared to acidic cell regions. The light response curve of NPQ shifted, after generation of an action potential, towards lower PFD. The release of NPQ by nigericin and the rapid reversal of action potential-triggered NPQ in darkness indicate its relation to thylakoid DeltapH. Generation of an action potential shortly after darkening converted the chloroplasts into a latent state with the F(m) identical to that of unexcited cells. This state transformed to the quenched state after turning on weak light that was insufficient for NPQ prior to membrane excitation of the cells. The ionophore, A23187, shifted NPQ plots similarly to the action potential effect, consistent with a likely role of a rise in the cytosolic Ca(2+) level in the action potential-induced quenching. The results suggest that a rapid electric signal, across the plasma membrane, might exert long-lived effects on photosynthesis and chlorophyll fluorescence through ion flux-mediated pathways.     

Fluorescence and Photosynthetic Activity of Chloroplasts in Acid and Alkaline Zones of Chara corallina

Continuous profiles of local pH near the cell surface of Chara corallinawere recorded during uniform longitudinal movement of an internodal cell relative to a stationary pH microelectrode. Under illumination, the pH profile consisted of alternating acid and alkaline bands with a pH difference of up to 3 pH units. After darkening, the bands disappeared and pH became uniformly distributed along the cell length. Chlorophyll fluorescence of chloroplasts was measured by microfluorometry at different locations within one cell, and significant differences were observed in close relation to light-dependent pH banding. The chlorophyll fluorescence yield was lower in zones of low external pH than in alkaline zones both under actinic and saturating light. The fluorescence parameters Fand F" _m and the quantum yield of photosystem II (PSII) displayed variations along the cell length in accordance with pH changes in unstirred layers of the medium. The results show that PSII photochemical efficiency and the rate of noncyclic electron transport are higher in the chloroplasts of acid zones (zones of H⁺extrusion from the cell) than in alkaline zones. The dependence of photosynthetic electron transport on local pH near the cell surface may result from different contents of CO₂in acid and alkaline regions. The acid zones are enriched with CO₂that readily permeates through the membrane providing the substrate for the Calvin cycle. Conversely, a poorly permeating form, HCO^– ₃is predominant in alkaline zones, which may restrict the dark reactions and photosynthetic electron flow.     

Effect of a Single Excitation Stimulus on Photosynthetic Activity and Light-dependent pH Banding in <Emphasis Type="Italic">Chara</Emphasis> Cells

Using pH microelectrodes and a Micro-scopy PAM (pulse-amplitude modulated) chlorophyll fluorometer, it is shown that a propagation of an action potential in Chara corallina leads to transient suppression of spatially periodic pH profiles along the illuminated cell. The suppression was manifested as a large pH decrease in the alkaline zones and a slight pH increase in the acid zones. The propagating action potential diminished the maximum yield of chlorophyll fluorescence (F_m′) in the alkaline cell regions, as well as the quantum yield of photosystem II photochemistry, without affecting F_m′ in the acid cell regions. The results indicate an interference of membrane excitation in the mechanisms responsible for pH banding patterns in Characean algae. Apparently, the electrical excitation of the plasma membrane in the alkaline cell regions initiates a pathway that can modulate membrane events at the thylakoid membrane.     

Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves

Higher plants must dissipate absorbed light energy that exceeds the photosynthetic capacity to avoid molecular damage to the pigments and proteins that comprise the photosynthetic apparatus. Described in this minireview is a current view of the biochemical, biophysical and bioenergetic aspects of the primary photoprotective mechanism responsible for dissipating excess excitation energy as heat from photosystem II (PSII). The photoprotective heat dissipation is measured as nonphotochemical quenching (NPQ) of the PSII chlorophyll a (Chl a) fluorescence. The NPQ mechanism is controlled by the trans-thylakoid membrane pH gradient (ΔpH) and the special xanthophyll cycle pigments. In the NPQ mechanism, the de-epoxidized endgroup moieties and the trans-thylakoid membrane orientations of antheraxanthin (A) and zeaxanthin (Z) strongly affect their interactions with protonated chlorophyll binding proteins (CPs) of the PSII inner antenna. The CP protonation sites and steps are influenced by proton domains sequestered within the proteo-lipid core of the thylakoid membrane. Xanthophyll cycle enrichment around the CPs may explain why changes in the peripheral PSII antenna size do not necessarily affect either the concentration of the xanthophyll cycle pigments on a per PSII unit basis or the NPQ mechanism. Recent time-resolved PSII Chi a fluorescence studies suggest the NPQ mechanism switches PSII units to an increased rate constant of heat dissipation in a series of steps that include xanthophyll de-epoxidation, CP-protonation and binding of the xanthophylls to the protonated CPs; the concerted process can be described with a simple two-step, pH-activation model. The xanthophyll cycle-dependent NPQ mechanism is profoundly influenced by temperatures suboptimal for photosynthesis via their effects on the trans-thylakoid membrane energy coupling system. Further, low temperature effects can be grouped into either short term (minutes to hours) or long term (days to seasonal) series of changes in the content and composition of the PSII pigment-proteins. This minireview concludes by briefly highlighting primary areas of future research interest regarding the NPQ mechanism.     

Pigment composition and functional state of the thylakoid membranes during preparation of samples for pigment-protein complexes separation by nondenaturing gel electrophoresis

The present study was conducted to examine changes in photosynthetic pigment composition and functional state of the thylakoid membranes during the individual steps of preparation of samples that are intended for a separation of pigmentprotein complexes by nondenaturing polyacrylamide gel electrophoresis. The thylakoid membranes were isolated from barley leaves (Hordeum vulgare L.) grown under low irradiance (50 μmol m⁻² s⁻¹). Functional state of the thylakoid membrane preparations was evaluated by determination of the maximal photochemical efficiency of photosystem (PS) II (F_V/F_M) and by analysis of excitation and emission spectra of chlorophyll a (Chl a) fluorescence at 77 K. All measurements were done at three phases of preparation of the samples: (1) in the suspensions of osmotically-shocked broken chloroplasts, (2) thylakoid membranes in extraction buffer containing Tris, glycine, and glycerol and (3) thylakoid membranes solubilized with a detergent decyl-β-D-maltosid. F_V/F_M was reduced from 0.815 in the first step to 0.723 in the second step and to values close to zero in solubilized membranes. Pigment composition was not pronouncedly changed during preparation of the thylakoid membrane samples. Isolation of thylakoid membranes affected the efficiency of excitation energy transfer within PSII complexes only slightly. Emission and excitation fluorescence spectra of the solubilized membranes resemble spectra of trimers of PSII light-harvesting complexes (LHCII). Despite a disrupted excitation energy transfer from LHCII to PSII antenna core in solubilized membranes, energy transfer from Chl b and carotenoids to emission forms of Chl a within LHCII trimers remained effective.     

Simulation of chlorophyll fluorescence rise and decay kinetics,and P<Subscript>700</Subscript>-related absorbance changes by using a rule-based kinetic Monte-Carlo method

A model of primary photosynthetic reactions in the thylakoid membrane was developed and its validity was tested by simulating three types of experimental kinetic curves: (1) the light-induced chlorophyll a fluorescence rise (OJIP transients) reflecting the stepwise transition of the photosynthetic electron transport chain from the oxidized to the fully reduced state; (2) the dark relaxation of the flash-induced fluorescence yield attributed to the Q_A^? oxidation kinetics in PSII; and (3) the light-induced absorbance changes near 820 or 705 nm assigned to the redox transitions of P₇₀₀ in PSI. A model was implemented by using a rule-based kinetic Monte-Carlo method and verified by simulating experimental curves under different treatments including photosynthetic inhibitors, heat stress, anaerobic conditions, and very high light intensity.     

Light-Triggered Action Potentials and Changes in Quantum Efficiency of Photosystem II in <Emphasis Type="Italic">Anthoceros</Emphasis> Cells

Capillary microelectrodes and pulse amplitude-modulated microfluorometry were used to study light-triggered changes in cell membrane potential, chlorophyll fluorescence, and photochemical yield of PSII in chloroplasts of a hornwort Anthoceros sp. The action potential was generated by illuminating the plant sample for a few seconds. It was accompanied by a reversible decrease in quantum efficiency of PSII and by nonphotochemical quenching of fluorescence that continued as long as 10 min after the light stimulus. The presence of ammonium ions (2 mM) enhanced the amplitude and prolonged the duration of dark changes of fluorescence parameters in accordance with the reported increase in duration and amplitude of the light-triggered action potential in the presence of NH ₄ ⁺ . A rapid retardation of PSII activity within the first seconds of illumination was also evident from absorbance changes at 810 nm reflecting the redox conversions of chlorophyll P700. The PSII-dependent stage of reduction in the induction curves of P700 absorbance was strongly suppressed, and the amplitudes of signals induced by white and far-red light (717 nm) differed insignificantly. It is concluded that a short-term irradiation triggers the generation of ΔpH at the thylakoid membranes, which is accompanied by inhibition of the plasma membrane H⁺ pump and by reversible inactivation of PSII due to increased thermal dissipation of chlorophyll excitations.     

Effect of Chlamydomonas plastid terminal oxidase 1 expressed in tobacco on photosynthetic electron transfer

The plastid terminal oxidase PTOX is a plastohydroquinone:oxygen oxidoreductase that is important for carotenoid biosynthesis and plastid development. Its role in photosynthesis is controversially discussed. Under a number of abiotic stress conditions, the protein level of PTOX increases. PTOX is thought to act as a safety valve under high light protecting the photosynthetic apparatus against photodamage. However, transformants with high PTOX level were reported to suffer from photoinhibition. To analyze the effect of PTOX on the photosynthetic electron transport, tobacco expressing PTOX‐1 from Chlamydomonas reinhardtii (Cr‐PTOX1) was studied by chlorophyll fluorescence, thermoluminescence, P700 absorption kinetics and CO₂ assimilation. Cr‐PTOX1 was shown to compete very efficiently with the photosynthetic electron transport for PQH₂. High pressure liquid chromatography (HPLC) analysis confirmed that the PQ pool was highly oxidized in the transformant. Immunoblots showed that, in the wild‐type, PTOX was associated with the thylakoid membrane only at a relatively alkaline pH value while it was detached from the membrane at neutral pH. We present a model proposing that PTOX associates with the membrane and oxidizes PQH₂ only when the oxidation of PQH₂ by the cytochrome b₆f complex is limiting forward electron transport due to a high proton gradient across the thylakoid membrane.     

Transient removal of alkaline zones after excitation of Chara cells is associated with inactivation of high conductance in the plasmalemma

The action potential (AP) of excitable plant cells is a multifunctional physiological signal. Its generation in characean algae suppresses the pH banding for 15–30 min and enhances the heterogeneity of spatial distribution of photosynthetic activity. This suppression is largely due to the cessation of H⁺ influx (OH⁻ efflux) in the alkaline cell regions. Measurements of local pH and membrane conductance in individual space-clamped alkaline zones (small cell areas bathed in an isolated pool of external medium) showed that the AP generation is followed by the transient disappearance of alkaline zone in parallel with a large decrease in membrane conductance. These changes, specific to alkaline zones, were only observed under continuous illumination following a relaxation period of at least 15 min after previous excitation. The excitation of dark-adapted cells produced no conductance changes in the post-excitation period. The results indicate that the origin of alkaline zones in characean cells is not due to operation of electroneutral H⁺/HCO₃⁻ symport or OH⁻/HCO₃⁻ antiport. It is concluded that the membrane excitation is associated with inactivation of plasmalemma high conductance in the alkaline cell regions.Key words: Chara, membrane conductance, pH banding, action potential, alkaline cell regions, heterogeneity     

Effects of light quality on the chloroplastic ultrastructure and photosynthetic characteristics of cucumber seedlings

To understand how light quality influences plant photosynthesis, we investigated chloroplastic ultrastructure, chlorophyll fluorescence and photosynthetic parameters, Rubisco and chlorophyll content and photosynthesis-related genes expression in cucumber seedlings exposed to different light qualities: white, red, blue, yellow and green lights with the same photosynthetic photon flux density of 100 μmol m^?2 s^?1. The results revealed that plant growth, CO₂ assimilation rate and chlorophyll content were significantly reduced in the seedlings grown under red, blue, yellow and green lights as compared with those grown under white light, but each monochromatic light played its special role in regulating plant morphogenesis and photosynthesis. Seedling leaves were thickened and slightly curled; Rubisco biosynthesis, expression of the rca, rbcS and rbcL, the maximal photochemical efficiency of PSII (Fv/Fm) and quantum yield of PSII electron transport (Ф_PSII) were all increased in seedlings grown under blue light as compared with those grown under white light. Furthermore, the photosynthetic rate of seedlings grown under blue light was significantly increased, and leaf number and chlorophyll content of seedlings grown under red light were increased as compared with those exposed to other monochromatic lights. On the contrary, the seedlings grown under yellow and green lights were dwarf with the new leaves etiolated. Moreover, photosynthesis, Rubisco biosynthesis and relative gene expression were greatly decreased in seedlings grown under yellow and green light, but chloroplast structural features were less influenced. Interestingly, the Fv/Fm, Ф_PSII value and chlorophyll content of the seedlings grown under green light were much higher than those grown under yellow light.     

Weak red light plays an important role in awakening the photosynthetic machinery following desiccation in the subaerial cyanobacterium Nostoc flagelliforme

The subaerial cyanobacterium Nostoc flagelliforme can survive for years in the desiccated state and light exposure may stimulate photosynthetic recovery during rehydration. However, the influence of light quality on photosynthetic recovery and the underlying mechanism remain unresolved. Exposure of field collected N. flagelliforme to light intensity ≥2 μmol photons m⁻² s⁻¹ showed that the speed of photosystem II (PSII) recovery was in the following order: red > green > blue ≈ violet light. Decreasing the light intensity showed that weak red light stimulated PSII recovery during rehydration. The chlorophyll fluorescence transient and oxygen evolution activity indicated that the oxygen evolution complex (OEC) was the activated site triggered by weak red light. The damaged D1 protein accumulated in the thylakoid membrane during dehydration and is degraded and resynthesized during dark rehydration. PsbO interaction with the thylakoid membrane was induced by weak red light. Thus, weak red light plays an important role in triggering OEC photoactivation and the formation of functional PSII during rehydration. In its arid habitats, weak red light could stimulate the awakening of dormant N. flagelliforme after absorbing water from nighttime dew or rain to maximize growth during the early daylight hours of the dry season.     

Drought-induced changes in chlorophyll fluorescence,photosynthetic pigments,and thylakoid membrane proteins of Vigna radiata

The effects of drought on chlorophyll fluorescence characteristics of PSII, photosynthetic pigments, thylakoid membrane protein (D1), and proline content in different varieties of mung bean plants were studied. Drought stress inhibits PSII activity and induces alterations in D1 protein. We observed a greater decline in the effective quantum yield of PSII, electron transport rate, and saturating photosynthetically active photon flux density (PPFD_sat) under drought stress in var. Anand than var. K-851 and var. RMG 268. This may possibly be due to either downregulation of photosynthesis or photoinhibition process. Withholding irrigation resulted in gradual diminution in total Chl content at Day 4 of stress. HPLC analysis revealed that the quantity of β-carotene in stressed plants was always higher reaching maxima at Day 4. Photoinactivation of PSII in var. Anand includes the loss of the D1 protein, probably from greater photosynthetic damage caused by drought stress than the other two varieties.     

Modelling of light-induced chlorophyll <Emphasis Type="Italic">a</Emphasis> fluorescence rise (O-J-I-P transient) and changes in 820 nm-transmittance signal of photosynthesis

Theoretical modelling is often overlooked in photosynthesis research even if it can significantly help with understanding of explored system. A new model of light-induced photosynthetic reactions occurring in and around thylakoid membrane is introduced here and used for theoretical modelling of not only the light-induced chlorophyll (Chl) a fluorescence rise (FLR; the O-J-I-P transient), reflecting function of photosystem II (PSII), but also of the 820 nmtransmittance signal (I₈₂₀), reflecting function of photosystem I (PSI) and plastocyanin (PC), paralleling the FLR. Correctness of the model was verified by successful simulations of the FLR and I₈₂₀ signal as measured with the control (no treatment) sample but also as measured with 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone- (inhibits electron transport in cytochrome b ₆/f) and methylviologen- (accepts electrons from iron-sulphur cluster of PSI) treated samples and with the control sample upon different intensities of excitation light. From the simulations performed for the control sample, contribution of the oxidised donor of PSI, P700, and oxidised PC to the I₈₂₀ signal minimum (reflects maximal accumulations of the two components) was estimated to be 75% and 25%, respectively. Further in silico experiments showed that PC must be reduced in the dark, cyclic electron transport around PSI must be considered in the model and activation of ferredoxin-NADP⁺-oxidoreductase (FNR) also affects the FLR. Correct simulations of the FLR and I₈₂₀ signal demonstrate robustness of the model, confirm that the electron transport reactions occurring beyond PSII affect the shape of the FLR, and show usefulness and perspective of theoretical approach in studying of the light-induced photosynthetic reactions.     

The effect of acid rain on ultrastructure and functional parameters of photosynthetic apparatus in pea leaves

The ultrastructure and functional parameters of the photosynthetic apparatus in leaves of 14-day-old pea seedlings were studied in conditions of laboratory simulated acid rain (SAR). Pea seedlings were sprayed with an aqueous solution containing NaNO₃ (0.2 mM) and Na₂SO₄ (0.2 mM) (pH 5.6, a control variant), or with the same solution, which was acidified to pH 2.5 (acid variant). Functional characteristics were determined by chlorophyll fluorescence analysis. There was reduction in the efficiency of the photosynthetic electron transport by 25% accompanied by an increase in the quantum yield of thermal dissipation of excess light quanta by 85% without significant change in maximum quantum yield of PSII photochemistry (F_v/F_m). Ultrastructural changes in chloroplasts were revealed by transmission electron microscopy (TEM) 2 days after the SAR treatment of pea leaves. In this case, changes in the structure of the grana and heterogeneity of the thylakoids packing in the granum, namely, an increase in thylakoid intraspace widths and thickness of granal thylakoids compared to the control, were found. It was shown also that carbonic anhydrase activity was significantly inhibited in chloroplast preparations isolated from SAR-treated pea leaves. We hypothesize possible involvement of chloroplast carbonic anhydrase in thylakoid granal structure maintenance. The structural disturbances and the inhibition of photochemical activity of chloroplasts are possible consequences of the carbonic anhydrase inactivation by SAR treatment leading to violation of HCO₃ ^?–CO₂ equilibrium. The data obtained suggest that acid rains negatively affect the photosynthetic apparatus by disrupting the membrane system of the chloroplast.     

Negative feedback regulation is responsible for the non-linear modulation of photosynthetic activity in plants and cyanobacteria exposed to a dynamic light environment

Photosynthetic organisms exposed to a dynamic light environment exhibit complex transients of photosynthetic activities that are strongly dependent on the temporal pattern of the incident irradiance. In a harmonically modulated light of intensity I≈const.+sin(ωt), chlorophyll fluorescence response consists of a steady-state component, a component modulated with the angular frequency of the irradiance ω and several upper harmonic components (2ω, 3ω and higher). Our earlier reverse engineering analysis suggests that the non-linear response can be caused by a negative feedback regulation of photosynthesis. Here, we present experimental evidence that the negative feedback regulation of the energetic coupling between phycobilisome and Photosystem II (PSII) in the cyanobacterium Synechocystis sp. PCC6803 indeed results in the appearance of upper harmonic modes in the chlorophyll fluorescence emission. Dynamic changes in the coupling of the phycobilisome to PSII are not accompanied by corresponding antiparallel changes in the Photosystem I (PSI) excitation, suggesting a regulation limited to PSII. Strong upper harmonic modes were also found in the kinetics of the non-photochemical quenching (NPQ) of chlorophyll fluorescence, of the P700 redox state and of the CO₂ assimilation in tobacco (Nicotiana tabaccum) exposed to harmonically modulated light. They are ascribed to negative feedback regulation of the reactions of the Calvin-Benson cycle limiting the photosynthetic electron transport. We propose that the observed non-linear response of photosynthesis may also be relevant in a natural light environment that is modulated, e.g., by ocean waves, moving canopy or by varying cloud cover. Under controlled laboratory conditions, the non-linear photosynthetic response provides a new insight into dynamics of the regulatory processes.     

玉米C4光合叶不同部位解剖结构和光抑制特性的比较

以玉米第54i全展叶（C4光合叶）为材料,分别测定基部、中部和顶部的光合速率后,将叶片置于强光（2000μmol·m^2·s^-1）下处理3h和暗中恢复3h,再测定这3个部位在处理期间的叶绿素荧光参数变化;然后分别从叶片的基部、中部和顶部取样观察显微结构和超微结构,测定叶绿素含量。结果表明,3个部位光合速率和叶绿素含量的大小依次为：中部〉顶部〉基部。基部的维管束鞘细胞叶绿体数量少,体积小,排列无规律,类囊体膜有部分垛叠;中部和顶部维管束鞘细胞叶绿体数量多,体积大,大部分围绕维管束呈离心排列,类囊体膜垛叠消失。在强光下,基部、中部和顶部均发生光抑制,但光抑制程度不同,根据严重度依次为：基部〉顶部〉中部,3个部位在暗中的光抑制恢复能力依次为：中部〉顶部〉基部。与叶基部相比,叶中部在强光下能维持较高的电子传递效率（φEo）和较低的热耗散比率（φDo）。这表明,C4光合循环是保持较高电子传递效率、减轻光抑制的重要因子。     

Effects of exogenous nitric oxide on the photosynthetic characteristics of bamboo (<Emphasis Type="Italic">Indocalamus barbatus</Emphasis> McClure) seedlings under acid rain stress

The effects of the exogenous application of nitric oxide (NO, in the form of sodium nitroprusside, SNP) on the diurnal variation in photosynthesis, chlorophyll content, chlorophyll fluorescence, light response curve and the net assimilation of CO₂ against intercellular CO₂ concentration (A-Ci) curve parameters were investigated in the leaves of bamboo (Indocalamus barbatus McClure) exposed to simulated acid rain (SAR, pH 3.0) stress. According to the results of the diurnal variation in photosynthesis, foliar applications of 100–400 mg/L SNP effectively inhibited the decrease in net photosynthetic rate (Pn) as a result of non-stomatal factors, and mitigated midday depression under acid rain stress. The mitigating effect was most pronounced at 400 mg/L SNP. However, at higher concentrations of SNP (700 and 1000 mg/L), the mitigating effect became weak and even counterproductive. The results of the chlorophyll content, light response and A-Ci curve parameters suggested that the regulating role of NO against acid rain in the photosynthetic processes occurs through improving not only the efficiency of the light-harvesting and the activity of photosynthetic apparatus, but also the absorption of CO₂ and the availability of CO₂ for photosynthesis. The results of the chlorophyll fluorescence investigation further indicated that NO protected PSII activity from the damage of acid rain toxicity by enhancing the electron transport activity and photochemical efficiency, especially concerning the increase in the proportion of PSII open reaction centers. Furthermore, NO induced an increase in photorespiration (Rp), rather than an increase in non-photochemical quenching (NPQ), to dissipate the excessive excitation energy, which provided some protection to the photosynthetic apparatus under acid rain stress.     

Growth of Anacystis in the Presence of Thiosulphate and its Consequences for the Architecture of the Photosynthetic Apparatus

With the aim of obtaining information on the degree of flexibility maintained in cyanobacteria in context with their phylogenetic position, Anacystis was grown in the presence of thiosulphate, oxidized in a photosystem I (PSI) dependent reaction (K_M 7.4 × 10^?3 M thiosulfate). Besides DBMIB, only o-phenanthroline and p-hydroxymercuribenzoate blocked thiosulphate-dependent PSI activity to some extent; iodonitrothymol, DCMU and cyanide had no influence. Growth of Anacystis in the presence of thiosulphate induced a reorganization of the photosynthetic apparatus characterized by a shift in the PSII/PSI ratio in favor of PSI, comparable to low light conditions. Capability for oxygenic photosynthesis never completely disappeared; structural elements of PSII were retained in the membrane to a certain degree. The antenna pigment system signalled high light under conditions of thiosulphate oxidation as judged from the ratio of phycocyanin to chlorophyll. Besides a shift in the ratio of PSII to PSI components, the polypeptide pattern of thylakoids from thiosulphate grown cells shows several additional components compared to the controls and, moreover, higher concentrations of some polypeptides present in the controls, particularly a M_r 41000 polypeptide. The process of thiosulphate oxidation appears bound to the thylakoid membrane.