Far‐red light enhances photochemical efficiency in a wavelength‐dependent manner

Linear electron transport depends on balanced excitation of photosystem I and II. Far‐red light preferentially excites photosystem I (PSI) and can enhance the photosynthetic efficiency when combined with light that over‐excites photosystem II (PSII). The efficiency of different wavelengths of far‐red light exciting PSI was quantified by measuring the change in quantum yield of PSII (Φ_PSII) of lettuce (Lactuca sativa) under red/blue light with narrowband far‐red light added (from 678 to 752 nm, obtained using laser diodes). The Φ_PSII of lettuce increased with increasing wavelengths of added light from 678 to 703 nm, indicating longer wavelengths within this region are increasingly used more efficiently by PSI than by PSII. Adding 721 nm light resulted in similar Φ_PSII as adding 703 nm light, but Φ_PSII tended to decrease as wavelength increased from 721 to 731 nm, likely due to decreasing absorptance and low photon energy. Adding 752 nm light did not affect Φ_PSII. Leaf chlorophyll fluorescence light response measurements showed lettuce had higher Φ_PSII under halogen light (rich in far‐red) than under red/blue light (which over‐excites PSII). Far‐red light is more photosynthetically active than commonly believed, because of its synergistic interaction with light of shorter wavelengths.     

Light quality during growth of Tradescantia albiflora regulates photosystem stoichiometry, photosynthetic function and susceptibility to photoinhibition

Tradescantia albiflora (Kunth) was grown under two different light quality regimes of comparable light quantity: in red + far-red light absorbed mainly by photosystem I (PSI light) and yellow light absorbed mainly by photosystem II (PSII light). The composition, function and ultrastructure of chloroplasts, and photoinhibition of photosynthesis in the two types of leaves were compared. In contrast to regulation by light quantity (Chow et al. 1991. Physiol. Plant. 81: 175–182), light quality exerted an effect on the composition of pigment complexes, function and structure of chloroplasts in Tradescantia: PSII light-grown leaves had higher Chl a/b ratios, higher PSI concentrations, lower PSII/PSI reaction centre ratios and less extensive thylakoid stacking than PSI light-grown leaves. Light quality triggered modulations of chloroplast components, leading to a variation of photosynthetic characteristics. A larger proportion of primary quinone acceptor (Q_A) in PSI light-grown leaves was chemically reduced at any given irradiance. It was also observed that the quantum yield of PSII photochemistry was lower in PSI light-grown leaves. PSI light-grown leaves were more sensitive to photoinihibition and recovery was slower compared to PSII light-grown leaves, showing that the PSII reaction centre in PSI light-grown leaves was more easily impaired by photoinhibition. The increase in susceptibility of leaves to photoinhibition following blockage of chloroplast-encoded protein synthesis was greater in PSII light-grown leaves, showing that these leaves normally have a greater capacity for PSII repair. Inhibition of zeaxanthin formation by dithiothreitol slightly increased sensitivity to photoinhibition in both PSI and PSII light-grown leaves.     

Action spectra of photosystems II and I and quantum yield of photosynthesis in leaves in State 1

The spectral global quantum yield (Y_II, electrons/photons absorbed) of photosystem II (PSII) was measured in sunflower leaves in State 1 using monochromatic light. The global quantum yield of PSI (Y_I) was measured using low-intensity monochromatic light flashes and the associated transmittance change at 810 nm. The 810-nm signal change was calibrated based on the number of electrons generated by PSII during the flash (4 · O₂ evolution) which arrived at the PSI donor side after a delay of 2 ms. The intrinsic quantum yield of PSI (y_I, electrons per photon absorbed by PSI) was measured at 712 nm, where photon absorption by PSII was small. The results were used to resolve the individual spectra of the excitation partitioning coefficients between PSI (a_I) and PSII (a_II) in leaves. For comparison, pigment–protein complexes for PSII and PSI were isolated, separated by sucrose density ultracentrifugation, and their optical density was measured. A good correlation was obtained for the spectral excitation partitioning coefficients measured by these different methods. The intrinsic yield of PSI was high (y_I = 0.88), but it absorbed only about 1/3 of quanta; consequently, about 2/3 of quanta were absorbed by PSII, but processed with the low intrinsic yield y_II = 0.63. In PSII, the quantum yield of charge separation was 0.89 as detected by variable fluorescence F_v/F_m, but 29% of separated charges recombined (Laisk A, Eichelmann H and Oja V, Photosynth. Res. 113, 145–155). At wavelengths less than 580 nm about 30% of excitation is absorbed by pigments poorly connected to either photosystem, most likely carotenoids bound in pigment–protein complexes.     

Dephosphorylation of photosystem II core proteins is light-regulated in vivo.

A number of photosystem II (PSII)-associated proteins, including D1, D2, CP43 and LHCII, are phosphorylated post-translationally by a membrane-bound, redox-regulated kinase activity. In vitro studies have demonstrated that these proteins can be dephosphorylated by membrane-bound phosphatase activity, reportedly insensitive to light or redox control. We demonstrate here that the PSII core proteins, D1, D2 and CP43, undergo light-stimulated, linear electron-transport-independent dephosphorylation in vivo. The in vivo dephosphorylation of D1 was characterized further and shown to depend upon light intensity, and to occur throughout the visible light spectrum with characteristics most consistent with light absorption by chlorophyll. PSII core protein dephosphorylation in vivo was stimulated by photosystem I (PSI)-specific far-red light, and inhibited by 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone, an inhibitor of plastoquinol oxidation by the cytochrome b6f complex. Based on these findings, we propose that PSI excitation is involved in regulating dephosphorylation of PSII core proteins in vivo.     

Adjustment of leaf photosynthesis to shade in a natural canopy: reallocation of nitrogen

The present study was performed to investigate the adjustment of the constituents of the light and dark reactions of photosynthesis to the natural growth irradiance in the leaves of an overstorey species, Betula pendula Roth, a subcanopy species Tilia cordata P. Mill., and a herb Solidago virgaurea L. growing in a natural plant community in Järvselja, Estonia. Shoots were collected from the site and properties of individual leaves were measured in a laboratory, by applying a routine of kinetic gas exchange and optical measurements that revealed photosystem II (PSII), photosystem I (PSI), and cytochrome b₆f densities per leaf area and the distribution of excitation (or chlorophyll, Chl) between the two photosystems. In parallel, N, Chl and ribulose-bisphosphate carboxylase-oxygenase (Rubisco) content was measured from the same leaves. The amount of N in photosynthetic proteins was calculated from the measured contents of the components of the photosynthetic machinery. Non-photosynthetic N was found as the residual of the budget. Growth in shade resulted in the decrease of leaf dry mass to a half of the DW in sun leaves in each species, but the total variation, from the top to the bottom of the canopy, was larger. Through the whole cross-section of the canopy, leaf dry weight (DW) and Rubisco content per area decreased by a factor of four, N content by a factor of three, but Chl content only by a factor of 1.7. PSII density decreased by a factor of 1.9, but PSI density by a factor of 3.2. The density of PSI adjusted to shade to a greater extent than the density of PSII. In shade, the distribution of N between the components of the photosynthetic machinery was shifted toward light-harvesting proteins at the expense of Rubisco. Non-photosynthetic N decreased the most substantially, from 54% in the sun leaves of B. pendula to 11% in the shade leaves of T. cordata. It is concluded that the redistribution of N toward light-harvesting Chl proteins in shade is not sufficient to keep the excitation rate of a PSII centre invariant. Contrary to PSII, the density of PSI – the photosystem that is in immediate contact with the carbon assimilation system – shade-adjusts almost proportionally with the latter, whereas its Chl antenna correspondingly increases. Even under N deficiency, a likely condition in the natural plant community, a substantial part of N is stored in non-photosynthetic proteins under abundant irradiation, but much less under limiting irradiation. At least in trees the general sequence of down-regulation due to shade adjustment is the following: (1) non-protein cell structures and non-photosynthetic proteins; (2) carbon assimilation proteins; (3) light reaction centre proteins, first PSI; and (4) chlorophyll-binding proteins.     

Adaptation to shade of the light-harvesting apparatus in Silene dioica

Abstract. The physiological characteristics and photo-system composition of the photosynthetic apparatus of Silene dioica , a woodland plant, grown in sun and natural shade are examined. As expected, shade leaves exhibited lower chlorophyll a/b ratios, light saturated rates of CO₂ assimilation (A_sat), dark respiration (R_d,) and light compensation points ( Г ), with both sun and shade leaves having similar absorptances and quantum yields of CO₂ assimilation (φ). Shade leaves were able to utilize far-red light for electron transport and carbon assimilation and reach the compensation point. Sun leaves in far-red light had a rate of carbon assimilation equivalent to their dark respiration rate. Chlorophyll fluorescence kinetics from leaves at 77 K together with analyses of thylakoid contents of photosystems (PS) I and II and the light-harvesting cholorphyll a/b protein complex associated with PSII (LHCII) demonstrated that the antenna size of PSII was similar in thylakoids of sun and shade leaves, but shade leaves contained ca. 20% more PSII and ca. 12% less PSI complexes. The increased PSII/PSI ratio in shade leaves accounted for their ability to achieve the compensation point in far-red light. An important feature of photosynethic shade adaptation in S. dioica is an increase in the PSII/PSI ratio and not an increase in the antenna size of PSII. The adaptive response of sun leaves when placed in a shade environment was rapid and had a half-time of ca. 18h.     

Cyanobacterial Acclimation to Photosystem I or Photosystem II Light

The organization and function of the photochemical apparatus of Synechococcus 6301 was investigated in cells grown under yellow and red light regimes. Broadband yellow illumination is absorbed preferentially by the phycobilisome (PBS) whereas red light is absorbed primarily by the chlorophyll (Chl) pigment beds. Since PBSs are associated exclusively with photosystem II (PSII) and most of the Chl with photosystem I (PSI), it follows that yellow and red light regimes will create an imbalance of light absorption by the two photosystems. The cause and effect relationship between light quality and photosystem stoichiometry in Synechococcus was investigated. Cells grown under red light compensated for the excitation imbalance by synthesis/assembly of more PBS-PSII complexes resulting in high PSII/PSI = 0.71 and high bilin/Chl = 1.30. The adjustment of the photosystem stoichiometry in red light-grown cells was necessary and sufficient to establish an overall balanced absorption of red light by PSII and PSI. Cells grown under yellow light compensated for this excitation imbalance by assembly of more PSI complexes, resulting in low PSII/PSI = 0.27 and low bilin/Chl = 0.42. This adjustment of the photosystem stoichiometry in yellow light-grown cells was necessary but not quite sufficient to balance the absorption of yellow light by the PBS and the Chl pigment beds. A novel excitation quenching process was identified in yellow light-grown cells which dissipated approximately 40% of the PBS excitation, thus preventing over-excitation of PSII under yellow light conditions. It is hypothesized that State transitions in O₂ evolving photosynthetic organisms may serve as the signal for change in the stoichiometry of photochemical complexes in response to light quality conditions.     

Photosynthetic acclimation of Tradescantia albiflora to growth irradiance: Lack of adjustment of light-harvesting components and its consequences

The photosynthetic acclimation of Tradescantia albiflora (Kunth), a trailing ground species naturally occurring in the deep shade of rainforests, was studied in relation to growth irradiance (glasshouse; direct light and 1 to 4 layers of shade cloth, giving 100 to 1.4% relative growth irradiance). Contrary to other irradiance studies of higher plants grown in natural habitats or controlled light environments, the chlorophyll a/b ratios of Tradescantia leaves were low (∼2.2) and constant. Acclimation to growth irradiance caused no changes in the relative amounts of specific Chl-proteins or the numbers of photosystem I (PSI) and PSII reaction centres on a chlorophyll basis, indicating that the light-harvesting antenna sizes of PSII and PSI, as well as the photosystem stoichiometry, were independent of growth irradiance. However, the amount of cytochrome f and ATP synthase on a chlorophyll basis increased with increasing the relative growth irradiance from 1.4 to 35%, showing acclimation of electron transport and photophosphorylation capacity. The photosynthetic capacity and ribulose 1, 5-bisphosphate carboxylase (EC 4.1.1.39) activity also increased with increase of the growth irradiance to 35%. Beyond that, the inflexible PSII/PSI stoichiometry and shade-type photosystem II/light-harvesting units in Tradescaniia are a disadvantage for long-term exposure to high irradiance since the leaves are more prone to photoinhibition.     

Far-red light-regulated efficient energy transfer from phycobilisomes to photosystem I in the red microalga Galdieria sulphuraria and photosystems-related heterogeneity of phycobilisome population

Phycobilisomes (PBS) are the major photosynthetic antenna complexes in cyanobacteria and red algae. In the red microalga Galdieria sulphuraria, action spectra measured separately for photosynthetic activities of photosystem I (PSI) and photosystem II (PSII) demonstrate that PBS fraction attributed to PSI is more sensitive to stress conditions and upon nitrogen starvation disappears from the cell earlier than the fraction of PBS coupled to PSII. Preillumination of the cells by actinic far-red light primarily absorbed by PSI caused an increase in the amplitude of the PBS low-temperature fluorescence emission that was accompanied by the decrease in PBS region of the PSI 77 K fluorescence excitation spectrum. Under the same conditions, fluorescence excitation spectrum of PSII remained unchanged. The amplitude of P700 photooxidation in PBS-absorbed light at physiological temperature was found to match the fluorescence changes observed at 77 K. The far-red light adaptations were reversible within 2-5min. It is suggested that the short-term fluorescence alterations observed in far-red light are triggered by the redox state of P700 and correspond to the temporal detachment of the PBS antenna from the core complexes of PSI. Furthermore, the absence of any change in the 77 K fluorescence excitation cross-section of PSII suggests that light energy transfer from PBS to PSI in G. sulphuraria is direct and does not occur through PSII. Finally, a novel photoprotective role of PBS in red algae is discussed.     

铅胁迫对玉米幼苗叶片光系统功能及光合作用的影响

通过同时测定玉米叶片的叶绿素荧光快速诱导动力学曲线和对820 nm光的吸收、分析叶片的气体交换过程以及叶绿体活性氧清除关键酶的活性,研究了不同浓度的铅(Pb)胁迫对玉米光系统Ⅰ(PSⅠ)、光系统Ⅱ(PSⅡ)的光化学活性和光合作用的影响,并分析了Pb胁迫下两个光系统的相互关系.结果表明:铅胁迫显著抑制了玉米地上部分和地下部分的生长、降低了叶片光合色素含量、并通过非气孔因素限制了光合作用、导致过剩激发能的增加;铅胁迫显著抑制了超氧化物歧化酶(SOD)和抗坏血酸过氧化物酶(APX)的活性、伤害了PSII反应中心、PSII的受体侧和供体侧(放氧复合体)以及PSI光化学活性.     

Loss of the precise control of photosynthesis and increased yield of non‐radiative dissipation of excitation energy after mild heat treatment of barley leaves

The after effects of a short exposure of intact barley leaves to moderately elevated temperature (40°C, 5 min) on the induction transients and the irradiance dependencies of photosynthesis and chlorophyll fluorescence are presented. This mild heat treatment strongly reduced the oscillations in the rate of photosynthesis and in the yield of chlorophyll fluorescence. However, only a 25% irreversible inhibition of maximum photosynthetic capacity of photosystem II (PSII) measured by oxygen evolution was produced and the intrinsic quantum yield of PSII measured by the chlorophyll fluorescence ratio (F_m‐ F_o)/F_m decreased by only 15%. In contrast, the above treatment increased radiationless dissipation processes in PSII by a factor of two. In heat‐treated leaves, photosynthesis was not saturated even by strong light. Both ΔpH‐dependent quenching of excitons in PSII (including formation of zeaxanthin) and state 1/state 2 transition were found to be stimulated. Heat exposure enhanced the control of PSII activity by PSI, as evidenced by a significant increase in the quenching effect of far‐red light on the maximum yield of chlorophyll fluorescence. It was deduced that after mild heat treatment, the photosynthetic apparatus in leaves lacks the precise coordinating control of electron transport and carbon metabolism owing to the inability of PSII to support electron transport at a level adequate for carbon metabolism. This effect was not related to the small irreversible thermal damage to PSII, but was rather due to a significant increase in non‐photochemical quenching of excitation energy.     

缺铁对大豆叶片光合作用和光系统Ⅱ功能的影响

通过气体交换和叶绿素荧光测定研究了缺铁对大豆叶片碳同化和光系统Ⅱ的影响。缺铁条件下大豆光合速率(Pn)大幅下降;最大光化学效率(po)下降幅度较小;荧光诱导动力学曲线发生明显的变化,其中电子传递活性明显下降,K相(VK)相对荧光产量提高。缺铁大豆的天线转化效率(Fv'/Fm')、光化学猝灭系数(qP)和光系统Ⅱ实际光化学效率(ΦPSⅡ)降低,而非光化学猝灭(NPQ)则明显增加。此外,缺铁大豆的光后荧光上升增强。据此,认为铁缺乏伤害了光系统Ⅱ复合物供体侧和受体侧的电子传递;缺铁条件下光系统I环式电子传递的增强可能在维持激发能耗散和ATP供给方面起一定作用。     

Demonstration of Two Sites of Inhibition of Electron Transport by Protein Phosphorylation in Wheat Thylakoids

The effect of protein phosphorylation on electron transportactivities of thylakoids isolated from wheat leaves was investigated.Protein phosphorylation resulted in a reduction in the apparentquantum yield of whole chain and photosystem II (PSII) electrontransport but had no effect on photosystem I (PSI) activity.The affinity of the D1 reaction centre polypeptide of PSII tobind atrazine was diminished upon phosphorylation, however,this did not reduce the light-saturated rate of PSII electrontransport. Phosphorylation also produced an inhibition of thelight-saturated rate of electron transport from water or durohydroquinoneto methyl viologen with no similar effect being observed onthe light-saturated rate of either PSII or PSI alone. This suggeststhat phosphorylation produces an inhibition of electron transportat a site, possibly the cytochrome b₆/f complex, between PSIIand PSI. This inhibition of whole-chain electron transport wasalso observed for thylakoids isolated from leaves grown underintermittent light which were deficient in polypeptides belongingto the light-harvesting chlorophyll-protein complex associatedwith photosystem II (LHCII). Consequently, this phenomenon isnot associated with phosphorylation of LCHII polypeptides. Apossible role for cytochrome b₆/f complexes in the phosphorylation-inducedinhibition of whole chain electron transport is discussed. Key words: Electron transport, light harvesting, photosystem 2, protein phosphorylation, thylakoid membranes, wheat (Triticum aestivum)     

State transitions in the cyanobacterium Synechococcus elongatus 7942 involve reversible quenching of the photosystem II core

Cyanobacteria use chlorophyll and phycobiliproteins to harvest light. The resulting excitation energy is delivered to reaction centers (RCs), where photochemistry starts. The relative amounts of excitation energy arriving at the RCs of photosystem I (PSI) and II (PSII) depend on the spectral composition of the light. To balance the excitations in both photosystems, cyanobacteria perform state transitions to equilibrate the excitation energy. They go to state I if PSI is preferentially excited, for example after illumination with blue light (light I), and to state II after illumination with green-orange light (light II) or after dark adaptation. In this study, we performed 77-K time-resolved fluorescence spectroscopy on wild-type Synechococcus elongatus 7942 cells to measure how state transitions affect excitation energy transfer to PSI and PSII in different light conditions and to test the various models that have been proposed in literature. The time-resolved spectra show that the PSII core is quenched in state II and that this is not due to a change in excitation energy transfer from PSII to PSI (spill-over), either direct or indirect via phycobilisomes.     

Cyclic electron flow plays an important role in photoprotection for the resurrection plant <Emphasis Type="Italic">Paraboea</Emphasis><Emphasis Type="Italic">rufescens</Emphasis> under drought stress

Resurrection plants could survive severe drought stress, but the underlying mechanism for protecting their photosynthetic apparatus against drought stress is unclear. Cyclic electron flow (CEF) has been documented as a crucial mechanism for photoprotection in Arabidopsis and tobacco. We hypothesized that CEF plays an important role in protecting photosystem I (PSI) and photosystem II (PSII) against drought stress for resurrection plants. To address this hypothesis, the effects of mild drought stress on light energy distribution in PSII and P700 redox state were examined in a resurrection plant Paraboea rufescens. Cyclic electron flow was not activated below the photosynthetic photon flux density (PPFD) of 400 μmol m⁻² s⁻¹ in leaves without drought stress. However, CEF was activated under low light in leaves with mild drought stress, and the effective quantum yield of PSII significantly decreased. Meanwhile, non-photochemical quenching (NPQ) was significantly stimulated not only under high light but also under low light. Compared with the control, the fraction of overall P700 that cannot be oxidized in a given state (PSI acceptor side limitation) under high light was maintained at low level of 0.1 in leaves with water deficit, indicating that the over-reduction of the PSI acceptor side was prevented by the significant stimulation of CEF. Furthermore, methyl viologen could significantly increase the PSII photo-inhibition induced by high light compared with chloramphenicol. These results suggested that CEF is an important mechanism for protecting PSI and PSII from drought stress in resurrection plants.     

Resolution of the Photosystem I and Photosystem II contributions to chlorophyll fluorescence of intact leaves at room temperature

Green leaves illuminated with photosynthetically active light emit red fluorescence, whose time-dependent intensity variations reflect photosynthetic electron transport (the Kautsky effect). Usually, fluorescence variations are discussed by considering only the contribution of PSII-associated chlorophyll a, although it is known that the fluorescence of PSI-associated chlorophyll a also contributes to the total fluorescence [Aust. J. Plant Physiol. 22 (1995) 131]. Because the fluorescence emitted by each photosystem cannot be measured separately by selecting the emission wavelength in in vivo conditions, the contribution of PSI to total fluorescence at room temperature is still in ambiguity. By using a diode array detector, we measured fluorescence emission spectra corresponding to the minimal (F(O)) and maximal (F(M)) fluorescence states. We showed that the different shapes of these spectra were mainly due to a higher contribution of PSI chlorophylls in the F(O) spectrum. By exciting PSI preferentially, we recorded a reference PSI emission spectrum in the near far-red region. From the F(O) and F(M) spectra and from this PSI reference spectrum, we derived specific PSI and PSII emission spectra in both the F(O) and F(M) states. This enables to estimate true value of the relative variable fluorescence of PSII, which was underestimated in previous works. Accurate separation of PSI-PSII fluorescence emission spectra will also enable further investigations of the distribution of excitation energy between PSI and PSII under in vivo conditions.     

Reversible photoinhibition of unhardened and cold-acclimated spinach leaves at chilling temperatures

The photoinhibition of photosynthesis at chilling temperatures was investigated in cold-acclimated and unhardened (acclimated to +18° C) spinach (Spinacia oleracea L.) leaves. In unhardened leaves, reversible photoinhibition caused by exposure to moderate light at +4° C was based on reduced activity of photosystem (PS) II. This is shown by determination of quantum yield and capacity of electron transport in thylakoids isolated subsequent to photoinhibition and recovery treatments. The activity of PSII declined to approximately the same extent as the quantum yield of photosynthesis of photoinhibited leaves whereas PSI activity was only marginally affected. Leaves from plants acclimated to cold either in the field or in a growth chamber (+1° C), were considerably less susceptible to the light treatment. Only relatively high light levels led to photoinhibition, characterized by quenching of variable chlorophyll a fluorescence (F_V) and slight inhibition of PSII-driven electron transport. Fluorescence data obtained at 77 K indicated that the photoinhibition of cold-acclimated leaves (like that of the unhardened ones) was related to increased thermal energy dissipation. But in contrast to the unhardened leaves, 77 K fluorescence of cold-acclimated leaves did not reveal a relative increase of PSI excitation. High-light-treated, cold-acclimated leaves showed increased rates of dark respiration and a higher light compensation point. The photoinhibitory fluorescence quenching was fully reversible in low light levels both at +18° C and +4° C; the recovery was much faster than in unhardened leaves. Reversible photoinhibition is discussed as a protective mechanism against excess light based on transformation of PSII reaction centers to fluorescence quenchers.Abbreviations F_O initial fluorescence - F_M maximal fluorescence - F_V devariable fluorescence (f_m-f_o) - PFD photon flux density - PS photosystem - SD standard deviation The authors thank the Deutsche Forschungsgemeinschaft and the Academy of Finland for financial support.     

Interaction of phycobilisomes with photosystem II dimers and photosystem I monomers and trimers in the cyanobacterium Spirulina platensis.

Distribution of phycobilisomes between photosystem I (PSI) and photosystem II (PSII) complexes in the cyanobacterium Spirulina platensis has been studied by analysis of the action spectra of H2 and O2 photoevolution and by analysis of the 77 K fluorescence excitation and emission spectra of the photosystems. PSI monomers and trimers were spectrally discriminated in the cell by the unique 760 nm low-temperature fluorescence, emitted by the trimers under reductive conditions. The phycobilisome-specific 625 nm peak was observed in the action spectra of both PSI and PSII, as well as in the 77 K fluorescence excitation spectra for chlorophyll emission at 695 nm (PSII), 730 nm (PSI monomers), and 760 nm (PSI trimers). The contributions of phycobilisomes to the absorption, action, and excitation spectra were derived from the in vivo absorption coefficients of phycobiliproteins and of chlorophyll. Analyzing the sum of PSI and PSII action spectra against the absorption spectrum and estimating the P700:P680 reaction center ratio of 5.7 in Spirulina, we calculated that PSII contained only 5% of the total chlorophyll, while PSI carried the greatest part, about 95%. Quantitative analysis of the obtained data showed that about 20% of phycobilisomes in Spirulina cells are bound to PSII, while 60% of phycobilisomes transfer the energy to PSI trimers, and the remaining 20% are associated with PSI monomers. A relevant model of organization of phycobilisomes and chlorophyll pigment-protein complexes in Spirulina is proposed. It is suggested that phycobilisomes are connected with PSII dimers, PSI trimers, and coupled PSI monomers.     

Photosynthetic responses of sun- and shade-grown barley leaves to high light: is the lower PSII connectivity in shade leaves associated with protection against excess of light?

In this study, we have compared photosynthetic performance of barley leaves (Hordeum vulgare L.) grown under sun and shade light regimes during their entire growth period, under field conditions. Analyses were based on measurements of both slow and fast chlorophyll (Chl) a fluorescence kinetics, gas exchange, pigment composition; and of light incident on leaves during their growth. Both the shade and the sun barley leaves had similar Chl a/b and Chl/carotenoid ratios. The fluorescence induction analyses uncovered major functional differences between the sun and the shade leaves: lower connectivity among Photosystem II (PSII), decreased number of electron carriers, and limitations in electron transport between PSII and PSI in the shade leaves; but only low differences in the size of PSII antenna. We discuss the possible protective role of low connectivity between PSII units in shade leaves in keeping the excitation pressure at a lower, physiologically more acceptable level under high light conditions.     

The State of Photosystem I Complexes in the Central Regions of Pea Granal Thylakoids

Grana-core and grana-margin fragments were obtained from pea (Pisum sativum L.) thylakoids, and both fractions contained photosystem I (PSI) complexes. The yield of these fractions exhibited variations for the plants grown during various periods of the summer season. Low-temperature fluorescence spectra, excitation spectra of long-wave fluorescence, and P700 kinetic characteristics were recorded for these fractions. PSI complexes in central granal regions were associated with PSII and the light-harvesting complexes of PSII, which followed from the excitation spectra of long-wave fluorescence and the kinetic characteristics of P700 light oxidation and dark reduction. The characteristics of the margin regions were changed depending on the fraction yield. If the yield was low, marginal fragments contained mainly PSI complexes. When the yield increased, PSI associates with PSII appeared. A spatial distribution and state of PSI complexes in granal thylakoids are discussed as related to the size and composition of the light-harvesting antenna.