Digitonin-sensitive LHCII enlarges the antenna of Photosystem I in stroma lamellae of Arabidopsis thaliana after far-red and blue-light treatment

Light drives photosynthesis. In plants it is absorbed by light-harvesting antenna complexes associated with Photosystem I (PSI) and photosystem II (PSII). As PSI and PSII work in series, it is important that the excitation pressure on the two photosystems is balanced. When plants are exposed to illumination that overexcites PSII, a special pool of the major light-harvesting complex LHCII is phosphorylated and moves from PSII to PSI (state 2). If instead PSI is over-excited the LHCII complex is dephosphorylated and moves back to PSII (state 1). Recent findings have suggested that LHCII might also transfer energy to PSI in state 1. In this work we used a combination of biochemistry and (time-resolved) fluorescence spectroscopy to investigate the PSI antenna size in state 1 and state 2 for Arabidopsis thaliana. Our data shows that 0.7 ± 0.1 unphosphorylated LHCII trimers per PSI are present in the stroma lamellae of state-1 plants. Upon transition to state 2 the antenna size of PSI in the stroma membrane increases with phosphorylated LHCIIs to a total of 1.2 ± 0.1 LHCII trimers per PSI. Both phosphorylated and unphosphorylated LHCII function as highly efficient PSI antenna.     

CO2 response of cyclic electron flow around PSI (CEF-PSI) in tobacco leaves--relative electron fluxes through PSI and PSII determine the magnitude of non-photochemical quenching (NPQ) of Chl fluorescence

We hypothesized that cyclic electron flow around photosystem I (CEF-PSI) participates in the induction of non-photochemical quenching (NPQ) of chlorophyll (Chl) fluorescence when the rate of photosynthetic linear electron flow (LEF) is electron-acceptor limited. To test this hypothesis, the relationships among photosynthesis rate, electron fluxes through both PSI and PSII [Je(PSI) and Je(PSII)] and Chl fluorescence parameters were analyzed simultaneously in intact leaves of tobacco plants at several light intensities and partial pressures of ambient CO2 (Ca). At low light intensities, decreasing Ca lowered the photosynthesis rate, but Je(PSI) and Je(PSII) remained constant. Je(PSI) was larger than Je(PSII), indicating the existence of CEF-PSI. Increasing the light intensity enhanced photosynthesis and both Je(PSI) and Je (PSII). Je(PSI)/Je(PSII) also increased at high light and at high light and low Ca combined, showing a strong, positive relationship with NPQ of Chl fluorescence. These results indicated that CEF-PSI contributed to the dissipation of photon energy in excess of that consumed by photosynthesis by driving NPQ of Chl fluorescence. The main physiological function of CEF-PSI in photosynthesis of higher plants is discussed.     

Reversible changes in structure and function of photosynthetic apparatus of pea (Pisum sativum) leaves under drought stress

The effects of drought on photosynthesis have been extensively studied, whereas those on thylakoid organization are limited. We observed a significant decline in gas exchange parameters of pea (Pisum sativum) leaves under progressive drought stress. Chl a fluorescence kinetics revealed the reduction of photochemical efficiency of photosystem (PS)II and PSI. The non-photochemical quenching (NPQ) and the levels of PSII subunit PSBS increased. Furthermore, the light-harvesting complexes (LHCs) and some of the PSI and PSII core proteins were disassembled in drought conditions, whereas these complexes were reassociated during recovery. By contrast, the abundance of supercomplexes of PSII-LHCII and PSII dimer were reduced, whereas LHCII monomers increased following the change in the macro-organization of thylakoids. The stacks of thylakoids were loosely arranged in drought-affected plants, which could be attributed to changes in the supercomplexes of thylakoids. Severe drought stress caused a reduction of both LHCI and LHCII and a few reaction center proteins of PSI and PSII, indicating significant disorganization of the photosynthetic machinery. After 7 days of rewatering, plants recovered well, with restored chloroplast thylakoid structure and photosynthetic efficiency. The correlation of structural changes with leaf reactive oxygen species levels indicated that these changes were associated with the production of reactive oxygen species.     

外源NO对NaHCO3胁迫下黑麦草幼苗光合生理响应的调节

采用营养液砂培方法,研究了外源一氧化氮(NO)对100 mmol/L NaHCO3胁迫下黑麦草幼苗叶片叶绿素含量、光合气体交换和叶绿素荧光参数、光能分配及叶黄素循环的影响。结果表明:(1)外施60μmol/L NO供体硝普钠(SNP)显著缓解了NaHCO3胁迫下叶绿素含量、净光合速率(Pn)、气孔导度(Gs)和气孔限制值(Ls)的下降及胞间CO2浓度(Ci)的升高,提高了光系统Ⅱ(PSⅡ)的潜在活性(Fv/Fo)、最大光化学效率(Fv/Fm)、实际光化学效率(ΦPSⅡ)和光化学猝灭(qP),降低了初始荧光(Fo)和非光化学猝灭(NPQ)。(2)NaHCO3胁迫下,外施SNP显著抑制了天线转换效率(Fv’/Fm’)的下降,降低了光系统间激发能分配的不平衡性(β/α-1)和天线热耗散的比例(D),提高了吸收光能中用于光化学反应的比例(P),而对PSⅡ反应中心的过剩光能(Ex)无明显影响。(3)外施SNP显著降低了NaHCO3胁迫下叶黄素循环库(V+A+Z)下降和叶黄素循环脱环氧化状态(A+Z)/(V+A+Z)上升的幅度。但SNP对NaHCO3胁迫的缓解效应可被NO清除剂血红蛋白(Hb)部分或完全地逆转,SNP的分解产物NaNO2处理对NaHCO3胁迫无明显改善。表明外源NO可能通过提高光化学效率,缓解了碱胁迫引起的光抑制对光合机构的破坏,从而提高黑麦草的光合效率。     

外源一氧化氮对干旱胁迫下杨树光合作用的影响

NO是生物体中一种自由基分子,其NO对树木叶片光合作用的影响研究未见报道.本文研究了外源NO对杨树叶片水分状况、光合作用和抗氧化物酶活力的调节作用.不同浓度SNP处理对杨树叶片含水量具有显著影响,杨树叶片含水率随着SNP浓度的提高而增加.当SNP浓度增加到00μmol·L^-1后各处理杨树叶片含水率变化趋于稳定.外源NO能提高水分胁迫下杨树叶片的光合、原初光能转化率Fv/Fm、Fm/Fo和Fv/Fo等的比值.其效果随水分胁迫时间的延长而降低.与此对应的是,短时间水分处理(1 h)的杨树叶片SOD和POD抗氧化物酶的活性显著高于长时间(3h)水分胁迫处理.SNP能显著提高不同干旱时间处理组的POD活性,而对SOD活性影响不明显.同时,随SNP浓度的增加,POD和SOD活性呈现先升后降的趋势.因此,干旱胁迫可引起杨树叶片光合效率降低,出现氧化伤害症状,外源NO可诱导抗氧化物酶POD和SOD活性的升高,缓解原初光能转化率Fv/Fm、Fm/Fo和Fv/Fo等值的降低,从而延缓活性氧积累,减轻水分胁迫对杨树叶片光合作用的影响.     

Dissipation of excess energy triggered by blue light in cyanobacteria with CP43′ (isiA)

The chlorophyll-protein CP43′ (isiA gene) induced by stress conditions in cyanobacteria is shown to serve as an antenna for Photosystem II (PSII), in addition to its known role as an antenna for Photosystem I (PSI). At high light intensity, this antenna is converted to an efficient trap for chlorophyll excitations that protects system II from photo-inhibition. In contrast to the ‘energy-dependent non-photochemical quenching’ (NPQ) in chloroplasts, this photoprotective energy dissipation in cyanobacteria is triggered by blue light. The induction is proportional to light intensity. Induction and decay of the quenching exhibit the same large temperature-dependence.     

Response of the photosynthetic apparatus of cotton (Gossypium hirsutum) to the onset of drought stress under field conditions studied by gas-exchange analysis and chlorophyll fluorescence imaging.

The functioning of the photosynthetic apparatus of cotton (Gossypium hirsutum) grown during the onset of water limitation was studied by gas-exchange and chlorophyll fluorescence to better understand the adaptation mechanisms of the photosynthetic apparatus to drought conditions. For this, cotton was grown in the field in Central Asia under well-irrigated and moderately drought-stressed conditions. The light and CO(2) responses of photosynthesis (A(G)), stomatal conductance (g(s)) and various chlorophyll fluorescence parameters were determined simultaneously. Furthermore, chlorophyll fluorescence images were taken from leaves to study the spatial pattern of photosystem II (PSII) efficiency and non-photochemical quenching parameters. Under low and moderate light intensity, the onset of drought stress caused an increase in the operating quantum efficiency of PSII photochemistry (varphi(PSII)) which indicated increased photorespiration since photosynthesis was hardly affected by water limitation. The increase in varphi(PSII) was caused by an increase of the efficiency of open PSII reaction centers (F(v)'/F(m)') and by a decrease of the basal non-photochemical quenching (varphi(NO)). Using a chlorophyll fluorescence imaging system a low spatial heterogeneity of varphi(PSII) was revealed under both irrigation treatments. The increased rate of photorespiration in plants during the onset of drought stress can be seen as an acclimation process to avoid an over-excitation of PSII under more severe drought conditions.     

The mechanism of cyclic electron flow

Apart from the canonical light-driven linear electron flow (LEF) from water to CO₂, numerous regulatory and alternative electron transfer pathways exist in chloroplasts. One of them is the cyclic electron flow around Photosystem I (CEF), contributing to photoprotection of both Photosystem I and II (PSI, PSII) and supplying extra ATP to fix atmospheric carbon. Nonetheless, CEF remains an enigma in the field of functional photosynthesis as we lack understanding of its pathway. Here, we address the discrepancies between functional and genetic/biochemical data in the literature and formulate novel hypotheses about the pathway and regulation of CEF based on recent structural and kinetic information.     

Competition between linear and cyclic electron flow in plants deficient in Photosystem I

Photosynthetic electron transport can involve either a linear flow from water to NADP, via Photosystems (PS) II and I or a cyclic flow just involving PSI. Little is known about factors regulating the relative flow through each of these pathways. We have examined photosynthetic electron transport through each system in plants of Arabidopsis thaliana in which either the PSI-D1 or PSI-E1 subunits of PSI have been knocked out. In both cases, this results in an imbalance in the turnover of PSI and PSII, such that PSII electron transport is limited by PSI turnover. Phosphorylation of light-harvesting complex II (LHCII) and its migration to PSI is enhanced but only partially reversible and not sufficient to balance photosystem turnover. In spite of this, cyclic electron flow is able to compete efficiently with PSI across a range of conditions. In dark-adapted leaves, the efficiency of cyclic relative to linear flow induced by far-red light is increased, implying that the limiting step of cyclic flow lies in the re-injection of electrons into the electron transport chain. Illumination of leaves with white light resulted in transient induction of a significant non-photochemical quenching in knockout plants which is probably high energy state quenching induced by cyclic electron flow. At high light and at low CO(2), non-photochemical quenching was greater in the knockout plants than in the wildtype. Comparison of PSI and PSII turnover under such conditions suggested that this is generated by cyclic electron flow around PSI. We conclude that, when the concentration of PSI is limiting, cyclic electron flow is still able to compete effectively with linear flow to maintain a high DeltapH to regulate photosynthesis.     

Proteomics, pigment composition, and organization of thylakoid membranes in iron-deficient spinach leaves

The changes induced in the photosynthetic apparatus of spinach (Spinacia oleracea L.) seedlings exposed to iron deficiency shortly after germination were characterized with two proteomic approaches coupled with chlorophyll and xanthophyll analysis and in vivo measurements of photosynthesis. During the first 10 d of iron deficiency the concentrations of chlorophyll b and violaxanthin were greatly reduced, but all xanthophylls recovered after 13-17 d of iron deficiency, when both chlorophylls were negatively affected. No new protein was formed in iron-deficient leaves, and no protein disappeared altogether. Photosystem I (PSI) proteins were largely reduced, but the stoichiometry of the antenna composition of PSI was not compromised. On the contrary, PSII proteins were less affected by the stress, but the specific antennae Lhcb4 and Lhcb6, Lhcb2 and its isoform Lhcb1.1 were all reduced, while the concentration of Lhcb3 increased. A strong reduction in thylakoid bending and an altered distribution pattern for the reduced PSI and PSII complexes were observed microscopically in iron-deficient leaves. Supercomplex organization was also affected by the stress. The trimeric organization of Lhcb and the dimerization of Lhca were reduced, while monomerization of Lhcb increased. However, the trimerization of Lhcb was partially recovered after 13-17 d of iron deficiency. In iron-deficient leaves, photosynthesis was strongly inhibited at different light intensities, and a high de-epoxidation status of the xanthophylls was observed, in association with a strong impairment of photochemical efficiency and an increase of heat dissipation as monitored by the non-photochemical quenching of fluorescence. All these negative effects of iron deficiency were attenuated but not fully reversed after again supplying iron to iron-deficient leaves for 7-13 d. These results indicate that iron deficiency has a strong impact on the proteomic structure of spinach photosystems and suggest that, in higher plants, adaptive mechanisms common in lower organisms, which allow rapid changes of the photosystem structure to cope with iron stress, are absent. It is speculated that the observed changes in the monomer-trimer equilibrium of major PSII antennae, which is possibly the result of xanthophyll fluctuations, is a first adaptative adjustment to iron deficiency, and may eventually play a role in light dissipation mechanisms.     

Modulation of photosynthetic electron transport in the absence of terminal electron acceptors: characterization of the rbcL deletion mutant of tobacco

Tobacco rbcL deletion mutant, which lacks the key enzyme Rubisco for photosynthetic carbon assimilation, was characterized with respect to thylakoid functional properties and protein composition. The Delta rbcL plants showed an enhanced capacity for dissipation of light energy by non-photochemical quenching which was accompanied by low photochemical quenching and low overall photosynthetic electron transport rate. Flash-induced fluorescence relaxation and thermoluminescence measurements revealed a slow electron transfer and decreased redox gap between Q(A) and Q(B), whereas the donor side function of the Photosystem II (PSII) complex was not affected. The 77 K fluorescence emission spectrum of Delta rbcL plant thylakoids implied a presence of free light harvesting complexes. Mutant plants also had a low amount of photooxidisible P700 and an increased ratio of PSII to Photosystem I (PSI). On the other hand, an elevated level of plastid terminal oxidase and the lack of F0 'dark rise' in fluorescence measurements suggest an enhanced plastid terminal oxidase-mediated electron flow to O2 in Delta rbcL thylakoids. Modified electron transfer routes together with flexible dissipation of excitation energy through PSII probably have a crucial role in protection of PSI from irreversible protein damage in the Delta rbcL mutant under growth conditions. This protective capacity was rapidly exceeded in Delta rbcL mutant when the light level was elevated resulting in severe degradation of PSI complexes.     

Quenching of excited states of chlorophyll molecules in submembrane fractions of Photosystem I by exogenous quinones

The ability of three substituted quinones, 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone (DBMIB), 2,6-dichloro-p-benzoquinone (DCBQ), and tetramethyl-p-benzoquinone (duriquinone) to quench the excited states of chlorophyll (Chl) molecules in Photosystem I (PSI) was studied. Chl fluorescence emission measured with isolated PSI submembrane fractions was reduced following the addition of exogenous quinones. This quenching progressively increased with rising concentrations of the exogenous quinones according to the Stern-Volmer law. The values of Stern-Volmer quenching coefficients were found to be 3.28 x 10(5) M(-1) (DBMIB), 1.31 x 10(4) M(-1) (DCBQ), and 3.7 x 10(3) M(-1) (duroquinone). The relative quenching capacities of the various exogenous quinones in PSI thus strictly coincided to those found for the quenching of Fo level of Chl fluorescence in isolated thylakoids, which is emitted largely by Photosystem II (PSII) [Biochim. Biophys. Acta (2003) 1604, 115-123]. Quenching of Chl excited states in PSI submembrane fractions by exogenous quinones slowed down the rate of P700, primary electron donor of PSI, photooxidation measured at limiting actinic light irradiances thus revealing a reduced photochemical capacity of absorbed quanta. The possible involvement of non-photochemical quenching of excited Chl states by oxidized phylloquinones, electron acceptors of PSI, and oxidized plastoquinones, mobile electron carriers between PSII and the cytochrome b(6)/f complex, into the control of photochemical activity of PSI is discussed.     

Acclimation of mesophyll and bundle sheath chloroplasts of maize to different irradiances during growth

The regulation by light of the photosynthetic apparatus, and composition of light-harvesting complexes in mesophyll and bundle sheath chloroplasts was investigated in maize. Leaf chlorophyll content, level of plastoquinone, PSI and PSII activities and Lhc polypeptide compositions were determined in plants grown under high, moderate and low irradiances. Photochemical efficiency of PSII, photochemical fluorescence quenching and non-photochemical fluorescence quenching over a range of actinic irradiances were also determined, using chlorophyll a fluorescence analysis. Acclimation of plants to different light conditions caused marked changes in light-harvesting complexes, LHCI and LHCII, and antenna complexes were also reorganized in these types of chloroplasts. The level of LHCII increased in plants grown in low light, even in agranal bundle sheath chloroplasts where the amount of PSII was strongly reduced. Irradiance also affected LHCI complex and the number of structural polypeptides, in this complex, generally decreased in chloroplasts from plants grown under lower light. Surprisingly moderate and low irradiances during growth do not affect the light reaction and fluorescence parameters of plants but generated differences in composition of light-harvesting complexes in chloroplasts. On the other hand, the changes in photosynthetic apparatus in plants acclimated to high light, resulted in a higher efficiency of photosynthesis. Based on these observations we propose that light acclimation to high light in maize is tightly coordinated adjustment of light reaction components/activity in both mesophyll and bundle sheath chloroplasts. Acclimation is concerned with balancing light utilization and level of the content of LHC complexes differently in both types of chloroplasts.     

不同形态钙对高温逆境下番茄叶片光合作用的调控作

以‘辽园多丽’番茄幼苗为试材,在高温胁迫下喷施氯化钙(CaCl₂)、纳米钙(Nano-Ca)和糖醇钙(Manntiol-Ca),研究不同形态钙处理对高温逆境下番茄叶片光合作用的调控作用.结果表明: 不同形态的钙处理均可抑制高温逆境下叶片中叶绿素a和类胡萝卜素含量的下降;显著提高净光合速率(P_n),不同程度提高蒸腾速率(T_r)和气孔导度(g_s);降低PSII非调节性能量耗散的量子产量\[Y(NO)\]和由于受体侧限制引起的PSI非光化学能量耗散的量子产量\[Y(NA)\],提高PSII调节性能量耗散的量子产量\[Y(NPQ)\]和由于供体侧限制引起的PSI非光化学能量耗散的量子产量\[Y(ND)\];提高叶片中钙含量;Manntiol-Ca和Nano-Ca处理的总体效果优于CaCl₂处理.与CaCl₂相比,Manntiol-Ca和Nano-Ca是提高高温逆境下番茄叶片光合作用更有效的钙制剂.     

Dissipation of excess energy triggered by blue light in cyanobacteria with CP43' (isiA)

The chlorophyll-protein CP43' (isiA gene) induced by stress conditions in cyanobacteria is shown to serve as an antenna for Photosystem II (PSII), in addition to its known role as an antenna for Photosystem I (PSI). At high light intensity, this antenna is converted to an efficient trap for chlorophyll excitations that protects system II from photo-inhibition. In contrast to the 'energy-dependent non-photochemical quenching' (NPQ) in chloroplasts, this photoprotective energy dissipation in cyanobacteria is triggered by blue light. The induction is proportional to light intensity. Induction and decay of the quenching exhibit the same large temperature-dependence.     

Competition between linear and cyclic electron flow in plants deficient in Photosystem I

Photosynthetic electron transport can involve either a linear flow from water to NADP, via Photosystems (PS) II and I or a cyclic flow just involving PSI. Little is known about factors regulating the relative flow through each of these pathways. We have examined photosynthetic electron transport through each system in plants of Arabidopsis thaliana in which either the PSI-D1 or PSI-E1 subunits of PSI have been knocked out. In both cases, this results in an imbalance in the turnover of PSI and PSII, such that PSII electron transport is limited by PSI turnover. Phosphorylation of light-harvesting complex II (LHCII) and its migration to PSI is enhanced but only partially reversible and not sufficient to balance photosystem turnover. In spite of this, cyclic electron flow is able to compete efficiently with PSI across a range of conditions. In dark-adapted leaves, the efficiency of cyclic relative to linear flow induced by far-red light is increased, implying that the limiting step of cyclic flow lies in the re-injection of electrons into the electron transport chain. Illumination of leaves with white light resulted in transient induction of a significant non-photochemical quenching in knockout plants which is probably high energy state quenching induced by cyclic electron flow. At high light and at low CO₂, non-photochemical quenching was greater in the knockout plants than in the wildtype. Comparison of PSI and PSII turnover under such conditions suggested that this is generated by cyclic electron flow around PSI. We conclude that, when the concentration of PSI is limiting, cyclic electron flow is still able to compete effectively with linear flow to maintain a high ΔpH to regulate photosynthesis.     

Quantification of photosystem I and II in different parts of the thylakoid membrane from spinach

Electron paramagnetic resonance (EPR) was used to quantify Photosystem I (PSI) and PSII in vesicles originating from a series of well-defined but different domains of the thylakoid membrane in spinach prepared by non-detergent techniques. Thylakoids from spinach were fragmented by sonication and separated by aqueous polymer two-phase partitioning into vesicles originating from grana and stroma lamellae. The grana vesicles were further sonicated and separated into two vesicle preparations originating from the grana margins and the appressed domains of grana (the grana core), respectively. PSI and PSII were determined in the same samples from the maximal size of the EPR signal from P700⁺ and Y_D, respectively. The following PSI/PSII ratios were found: thylakoids, 1.13; grana vesicles, 0.43; grana core, 0.25; grana margins, 1.28; stroma lamellae 3.10. In a sub-fraction of the stroma lamellae, denoted Y-100, PSI was highly enriched and the PSI/PSII ratio was 13. The antenna size of the respective photosystems was calculated from the experimental data and the assumption that a PSII center in the stroma lamellae (PSIIβ) has an antenna size of 100 Chl. This gave the following results: PSI in grana margins (PSIα) 300, PSI (PSIβ) in stroma lamellae 214, PSII in grana core (PSIIα) 280. The results suggest that PSI in grana margins have two additional light-harvesting complex II (LHCII) trimers per reaction center compared to PSI in stroma lamellae, and that PSII in grana has four LHCII trimers per monomer compared to PSII in stroma lamellae. Calculation of the total chlorophyll associated with PSI and PSII, respectively, suggests that more chlorophyll (about 10%) is associated with PSI than with PSII.     

Upregulation of bundle sheath electron transport capacity under limiting light in C4 Setaria viridis

C₄ photosynthesis is a biochemical pathway that operates across mesophyll and bundle sheath (BS) cells to increase CO₂ concentration at the site of CO₂ fixation. C₄ plants benefit from high irradiance but their efficiency decreases under shade, causing a loss of productivity in crop canopies. We investigated shade acclimation responses of Setaria viridis, a model monocot of NADP-dependent malic enzyme subtype, focussing on cell-specific electron transport capacity. Plants grown under low light (LL) maintained CO₂ assimilation rates similar to high light plants but had an increased chlorophyll and light-harvesting-protein content, predominantly in BS cells. Photosystem II (PSII) protein abundance, oxygen-evolving activity and the PSII/PSI ratio were enhanced in LL BS cells, indicating a higher capacity for linear electron flow. Abundances of PSI, ATP synthase, Cytochrome b₆f and the chloroplast NAD(P)H dehydrogenase complex, which constitute the BS cyclic electron flow machinery, were also increased in LL plants. A decline in PEP carboxylase activity in mesophyll cells and a consequent shortage of reducing power in BS chloroplasts were associated with a more oxidised plastoquinone pool in LL plants and the formation of PSII – light-harvesting complex II supercomplexes with an increased oxygen evolution rate. Our results suggest that the supramolecular composition of PSII in BS cells is adjusted according to the redox state of the plastoquinone pool. This discovery contributes to the understanding of the acclimation of PSII activity in C₄ plants and will support the development of strategies for crop improvement, including the engineering of C₄ photosynthesis into C₃ plants.     

Photosystem II photoinhibition-repair cycle protects Photosystem I from irreversible damage

Photodamage of Photosystem II (PSII) has been considered as an unavoidable and harmful reaction that decreases plant productivity. PSII, however, has an efficient and dynamically regulated repair machinery, and the PSII activity becomes inhibited only when the rate of damage exceeds the rate of repair. The speed of repair is strictly regulated according to the energetic state in the chloroplast. In contrast to PSII, Photosystem I (PSI) is very rarely damaged, but when occurring, the damage is practically irreversible. While PSII damage is linearly dependent on light intensity, PSI gets damaged only when electron flow from PSII exceeds the capacity of PSI electron acceptors to cope with the electrons. When electron flow to PSI is limited, for example in the presence of DCMU, PSI is extremely tolerant against light stress. Proton gradient (ΔpH)-dependent slow-down of electron transfer from PSII to PSI, involving the PGR5 protein and the Cyt b6f complex, protects PSI from excess electrons upon sudden increase in light intensity. Here we provide evidence that in addition to the ΔpH-dependent control of electron transfer, the controlled photoinhibition of PSII is also able to protect PSI from permanent photodamage. We propose that regulation of PSII photoinhibition is the ultimate regulator of the photosynthetic electron transfer chain and provides a photoprotection mechanism against formation of reactive oxygen species and photodamage in PSI.     

Effect of the anthocyanic epidermal layer on Photosystem II and I energy dissipation processes in Tradescantia pallida (Rose) Hunt

The effect of anthocyanic cells of the epidermal layer was investigated on photosynthetic activity of the higher plant Tradescantia pallida. To determine the possible indirect role of anthocyanin in photosynthesis, analysis was done on intact leaves and leaves where anthocyanic epidermal layer was removed. Energy dissipation processes related to Photosystem II (PSII) and Photosystem I (PSI) activity was done using simultaneously Chlorophyll a (Chl a) fluorescence and P700 transmittance signals change. In anthocyanic epidermal-less leaves, PSII photochemical activity was more decreased in dependence to increasing light irradiance exposure. We found that photoinhibition of PSII decreased PSI activity by reducing the electron flow toward PSI, especially under high light intensities. Under those conditions, it resulted in the accumulation of oxidized PSI reaction centers, which was stronger in leaves where the anthocyanic epidermal layer was removed. In conclusion, our results showed that the anthocyanic epidermal layer had a photoprotective effect only on the PSII and not on the PSI of T. pallida leaves, supporting the role of anthocyanin pigments in the regulation of photosynthesis for excess absorbed light irradiance.