Photosynthetic efficiency during ontogenesis of leaves of Betula pendula

Abstract. Net photosynthesis, photosynthetic electron transport, and leaf area density of photosynthetic units have been studied in developing, mature, and old leaves of seedlings of Betula pendula . The photosynthetic quantum yield under light-limiting conditions and the leaf area related rale of light-saturated net photosynthesis were lower in developing than in mature and old leaves. Developing leaves also had more oxygen inhibition of photosynthesis, a lower pool size of plastoquinone in the electron transport chain, a lower chlorophyll content and a lower leaf area density of photosynthetic units than mature and old leaves. The photosynthetic properties of The oldest leaves resulted partly from acclimation to shade and partly from a different ontogeny to that of younger leaves.     

Formation and development of photosynthetic units in repigmenting Rhodopseudomonas sphaeroides wild type and "Phofil" mutant strain.

The formation of the photosynthetic apparatus in the wild type Rhodopseudomonas sphaeroides and in the "Phofil" mutant was investigated by analyzing absorption and fluorescence parameters and the kinetics of fluorescence induction. Repigmentation was induced by transfer of bleached cells to semi-aerobic culture conditions. 1. In the "Phofil" mutant, functional photosynthetic units appear at pigment cellular contents which depend on the physiological state of the inoculum. The unadapted mutant can reach the functional units stage at a cellular pigment level 20 times lower than that of the wild type, although the size and composition of the units are identical in both strains. This rules out the hypothesis of photosynthetic units forming by random integration of pigments into the membrane. Moreover, the fact that units are separate at this stage (as shown by the exponential shape of fluorescence induction kinetics) suggests that the units' formation proceeds from discrete predetermined membrane sites. 2. In repigmentng wild type cells, the multistep assembly of bacteriochlorophyll complexes appears correlated to their organization in photosynthetic units as follows: (i) During a first stage, B-875 is mostly synthesized, while the efficiency of transfer increases, suggesting that the pigments are initially interpersed at comparatively large average distances from each other in the bleached membrane. (ii) After 1.5 h of repigmentation, the transfer and trapping efficiencies reach a maximum. The units (26 B-875 + 11 B-850 per center) are still separate, as shown by exponential fluorescence kinetics. (iii) The increase in the units' size then essentially proceeds through B-850 incorporation. Energy transfer between units occurs at a comparatively late stage, i.e., 5 h repigmentation.     

Photosynthetic productivity of conical helical tubular photobioreactors incorporating Chlorella sp. under various culture medium flow conditions

The characteristics of the flow of culture medium significantly affects the photosynthetic productivity of bioreactors incorporating microalgae. Therefore, in order to optimize the performance of a conical helical tubular photobioreactor (CHTP) designed to be useful in practical applications, we characterized the flow pattern of the culture medium through the reactor. The effects of medium flow conditions on the photosynthetic productivity of Chlorella sp. were investigated using several different CHTP units with 0.50-m2 installation areas which were designed to vary the direction and rate of flow driven by airlift. In addition, the performance of two- and four-unit systems constructed by combining individual CHTP units was evaluated. We found that when medium flowed from the bottom to the top of the photostage, it exhibited smoother flow of culture medium than when flowing from top to bottom, which led to higher photosynthetic productivity by the former. Consistent with theoretical calculations, varying the lengths of vertical flow passages caused flow rates to vary, and higher flow rates meant smoother circulation of medium and better photosynthetic performance. Flow of medium through a four-unit CHTP system was similar to that in single units, enabling a photosynthetic productivity of 31.0 g-dry biomass per m2-installation area per day to be achieved, which corresponded to a photosynthetic efficiency of 7.50% (photosynthetically active radiation (PAR; 400-700 nm)). This high photosynthetic performance was possible because smoother medium flow attained in single units was also attained in the four-unit system.     

Photosynthesis in trees: organization of chlorophyll and photosynthetic unit size in isolated gymnosperm chloroplasts

Chloroplasts have been isolated in high yield from several gymnosperms and from two deciduous trees. The organization of chlorophyll in the chloroplasts of these woody species is basically similar to that in angiosperm crop plants and green algae. The tree chloroplasts contain two chlorophyll proteins, the P700-chlorophyll a-protein and the major light-harvesting chlorophyll a/b-protein, the size, spectral characteristics, and function of which are the same as the equivalent complexes previously isolated from other classes of green plants. All the gymnosperms have chlorophyll/P700 ratios (photosynthetic unit sizes) 1.6 to 3.8 times larger than that typically found in crop plants; the deciduous trees have units of intermediary size. The presence of fewer but larger photosynthetic units in the woody species can partially account for their lower photosynthetic rate and explains why their photosynthetic processes saturate at lower light intensities. Chloroplasts of shade needles have large units containing a greater proportion of the light-harvesting chlorophyll a/b-protein than those of sun needles.     

Construction of hybrid photosynthetic units using peripheral and core antennae from two different species of photosynthetic bacteria: detection of the energy transfer from bacteriochlorophyll a in LH2 to bacteriochlorophyll b in LH1

Typical purple bacterial photosynthetic units consist of supra-molecular arrays of peripheral (LH2) and core (LH1-RC) antenna complexes. Recent atomic force microscopy pictures of photosynthetic units in intact membranes have revealed that the architecture of these units is variable (Scheuring et al. (2005) Biochim Bhiophys Acta 1712:109–127). In this study, we describe methods for the construction of heterologous photosynthetic units in lipid-bilayers from mixtures of purified LH2 (from Rhodopseudomonas acidophila) and LH1-RC (from Rhodopseudomonas viridis) core complexes. The architecture of these reconstituted photosynthetic units can be varied by controlling ratio of added LH2 to core complexes. The arrangement of the complexes was visualized by electron-microscopy in combination with Fourier analysis. The regular trigonal array of the core complexes seen in the native photosynthetic membrane could be regenerated in the reconstituted membranes by temperature cycling. In the presence of added LH2 complexes, this trigonal symmetry was replaced with orthorhombic symmetry. The small lattice lengths for the latter suggest that the constituent unit of the orthorhombic lattice is the LH2. Fluorescence and fluorescence-excitation spectroscopy was applied to the set of the reconstituted membranes prepared with various proportions of LH2 to core complexes. Remarkably, even though the LH2 complexes contain bacteriochlorophyll a, and the core complexes contain bacteriochlorophyll b, it was possible to demonstrate energy transfer from LH2 to the core complexes. These experiments provide a first step along the path toward investigating how changing the architecture of purple bacterial photosynthetic units affects the overall efficiency of light-harvesting.     

Light-Shade Adaptation : TWO STRATEGIES IN MARINE PHYTOPLANKTON

Using chlorophyll/P700 ratios, the size and number of photosynthetic units were estimated, as a function of light-shade adaptation in two species of marine phytoplankton: Skeletonema costatum, a diatom, and Dunaliella tertiolecta, a chlorophyte. In the diatom, light-shade adaptation is characterized primarily by changes in the size and not the number of P700 units, whereas in the chlorophyte, overall changes in chlorophyll content are related to changes in the number and not the size of P700 units. A correlation between the characteristics of P700 units and photosynthetic responses was not established. Both strategies of light-shade adaptation effectively harvest and transfer light energy to reaction centers, however, the Skeletonema strategy is more effective at subsaturating intensities. The two strategies may represent an evolutionary divergence in photosynthetic adaptation to variations in light intensity.     

Energy transfer in the photosynthetic unit. I. The concept of independent units for photosystem II analyzed by flash yields for dichlorophenolindophenol reduction

The problem of the interconnection of photosynthetic units is dealt with. Flash yield results together with quantum yield measurements of DCPIP reduction in isolated chloroplasis indicate that photosynthetic units of PS-II are essentially independent and probably morphological entities. This is shown by the exponential dependence of the flash yield on the flash intensity and a high quantum yield of excitation trapping. Deviation from exponentiality observed in some samples is explained in terms of energy transfer between the units when they are densely packed.     

Sequential assembly of photosynthetic units in Rhodobacter sphaeroides as revealed by fast repetition rate analysis of variable bacteriochlorophyll a fluorescence

The development of functional photosynthetic units in Rhodobacter sphaeroides was followed by near infra-red fast repetition rate (IRFRR) fluorescence measurements that were correlated to absorption spectroscopy, electron microscopy and pigment analyses. To induce the formation of intracytoplasmic membranes (ICM) (greening), cells grown aerobically both in batch culture and in a carbon-limited chemostat were transferred to semiaerobic conditions. In both aerobic cultures, a low level of photosynthetic complexes was observed, which were composed of the reaction center and the LH1 core antenna. Interestingly, in the batch cultures the reaction centers were essentially inactive in forward electron transfer and exhibited low photochemical yields F(V)/F(M), whereas the chemostat culture displayed functional reaction centers with a rather rapid (1-2 ms) electron transfer turnover, as well as a high F(V)/F(M) of approximately 0.8. In both cases, the transfer to semiaerobiosis resulted in rapid induction of bacteriochlorophyll a synthesis that was reflected by both an increase in the number of LH1-reaction center and peripheral LH2 antenna complexes. These studies establish that photosynthetic units are assembled in a sequential manner, where the appearance of the LH1-reaction center cores is followed by the activation of functional electron transfer, and finally by the accumulation of the LH2 complexes.     

Photosynthesis in relation to leaf characteristics of cotton from controlled and field environments

In situ and light-saturated net photosynthetic rates per unit leaf area were greater in cotton (Gossypium hirsutum L.) plants grown in pots in the field than in similar plants from a phytotron growth chamber. Light-saturated stomatal resistances did not differ in leaves of similar age and exposure on field and chamber plants; lower photosynthetic rates in chamber leaves were associated with greater mesophyll resistance. Differences in net photosynthetic rates were related to differences in leaf thickness. When the photosynthetic rates were expressed per unit of mesophyll volume or per unit chlorophyll differences between field and chamber plants were much less than when rates were expressed per unit leaf area. Characterization of the chloroplast lamellar proteins showed that the field leaves had smaller photosynthetic units than the chamber leaves. Since the field leaves also contained more chlorophyll per unit area, this resulted in a much larger number of photosynthetic units per unit area in the field leaves.     

Analysis of picosecond laser induced fluorescence phenomena in photosynthetic membranes utilizing a master equation approach.

A Pauli master equation is formulated and solved to describe the fluorescence quantum yield, phi, and the fluorescence temporal decay curves. F(t), obtained in picosecond laser excitation experiments of photosynthetic systems. It is assumed that the lowering of phi with increasing pulse intensity is due to bimolecular singlet exciton annihilation processes which compete with the monomolecular exciton decay processes; Poisson statistics are taken into account. Calculated curves of phi as a function of the number of photon hits per domain are compared with experimental data, and it is concluded that these domains contain at least two to four connected photosynthetic units (depending on the temperature), where each photosynthetic unit is assumed to contain approximately 300 pigment molecules. It is shown that under conditions of high excitation intensities, the fluorescence decays approximately according to the (time)1/2 law.     

Photosynthetic Units

Leaf tissues of aurea mutants of tobacco and Lespedeza have been shown to have higher photosynthetic capacity per molecule of chlorophyll, a higher saturation intensity, a simpler lamellar structure, and the same quantum yield as their dark green parents. Here we report on the values of photosynthetic units for both types of plants and some algae. The unit has been assumed to be about as uniform and steady in the plant world as the quantum efficiency. The number on which all theoretical discussions have been based so far is 2400 per O₂ evolved or CO₂ reduced. With dark green plants and algae our determinations of units by means of 40 µsec flashes superimposed on a steady rate of background photosynthesis at 900 ergs cm^-2 sec^-1 of red light yielded mostly numbers between 2000 and 2700. However, the photosynthetic unit turned out to be very variable, even in these objects. In aurea mutants the unit was distinctly smaller, averaging 600 chl/CO₂. By choosing the right combination of colors for flash and background light, units as low as 300 chl/CO₂ or 40 chl/e^- could be measured consistently. We found five well-defined groups of units composed of multiples of its smallest member. These new findings are discussed in terms of structural entities that double or divide under the influence of far-red light.     

Frost hardening and photosynthetic performance of Scots pine (Pinus sylvestris L.) needles. I. Seasonal changes in the photosynthetic apparatus and its function

Photosynthetic CO₂ uptake, the photochemical efficiency of photosystem II, the contents of chlorophyll and chlorophyll-binding proteins, and the degree of frost hardiness were determined in three-year-old Scots pine (Pinus sylvestris L.) trees growing in the open air but under controlled daylength. The following conditions were compared: 9-h light period (short day), 16-h light period (long day), and natural daylength. Irrespective of induction by short-day photoperiods or by subfreezing temperatures, frost hardening of the trees was accompanied by a long-lasting pronounced decrease in the photosynthetic rates of one-year-old needles. Under moderate winter conditions, trees adapted to a long-day photoperiod, assimilated CO₂ with higher rates than the short-day-treated trees. In the absence of strong frost, photochemical efficiency was lower under short-day conditions than under a long-day photoperiod. Under the impact of strong frost, photochemical efficiency was strongly inhibited in both sets of plants. The reduction in photosynthetic performance during winter was accompanied by a pronounced decrease in the content of chlorophyll and of several chlorophyll-binding proteins [light-harvesting complex (LHC)IIb, LHC Ib, and a chlorophyll-binding protein with MW 43 kDa (CP 43)]. This observed seasonal decrease in photosynthetic pigments and in pigment-binding proteins was irrespective of the degree of frost hardiness and was apparantly under the control of the length of the daily photoperiod. Under a constant 9-h daily photoperiod the chlorophyll content of the needles was considerably lower than under long-day conditions. Transfer of the trees from short-day to long-day conditions resulted in a significantly increased chlorophyll content, whereas the chlorophyll content decreased when trees were transferred from a long-day to a short-day photoperiod. The observed changes in photosynthetic pigments and pigment-binding proteins in Scots pine needles are interpreted as a reduction in the number of photosynthetic units induced by shortening of the daily light period during autumn. This results in a reduction in the absorbing capacity during the frost-hardened state. Received: 3 March 1997 / Accepted: 16 July 1997     

Capture frequency of excitations and energy transfer between photosynthetic units in the photosystem II

The connections which exist, in the photosynthetic apparatus, between the spatial arrangement of chlorophyll and the movement of excitation energy are discussed.The capture frequency of excitations in the photosystem II is analysed. At the microscopic level of a photosynthetic unit two stages are studied: the propagation of the excitation to the reaction centre and the photochemical utilization of the excitation by the centre. It is shown that the transport process is not a limiting one. It implies that the capture frequency depends on the reaction centre state. Thus it is possible to distinguish eight states for a photosynthetic unit of system II.At the macroscopic level of a set of units, the analysis of the fluorescence yield-fluctuations shows that these units are not isolated. It also indicates that the fluorescence emitted by the photosynthetic apparatus originates almost entirely from the system II, and that the reaction centres are traps for excitations whatever their states.     

基因转录介导同质园条件下拟南芥不同地理种群的光合能力分化

不同环境条件决定着植物光合能力的多态性,但在相同环境条件下同种植物不同种群间表现出光合能力分化的内在机制仍不清楚,本文旨在揭示同质园条件下欧洲拟南芥(Arabidopsis thaliana)不同地理种群光合能力的分化以及其基因转录调控机制。在同质园条件下,测定来自欧洲不同气候区的23个拟南芥的地理种群的气体交换参数、叶绿素荧光参数及SPAD值综合比较其光合能力差异。另外,根据测定结果选取光合能力差异的典型种群,利用实时定量PCR技术对光合相关基因表达水平进行验证。比较研究发现欧洲不同气候区拟南芥的地理种群间的气体交换参数差异较大,其中净光合速率的变化范围为2～11μmol·m^-2·s^-1;而叶绿素荧光参数变异幅度较小,变异幅度几乎不超过10%。聚类分析表明23个拟南芥种群被分为强光合能力和弱光合能力2组,强光合种群主要分布在中欧和西欧地区,净光合速率平均为7.37μmol·m^-2·s^-1;而弱光合种群则主要分布在东欧和南欧,净光合速率平均为4.46μmol·m^-2·s     

Sequential assembly of photosynthetic units in Rhodobacter sphaeroides as revealed by fast repetition rate analysis of variable bacteriochlorophyll a fluorescence

The development of functional photosynthetic units in Rhodobacter sphaeroides was followed by near infra-red fast repetition rate (IRFRR) fluorescence measurements that were correlated to absorption spectroscopy, electron microscopy and pigment analyses. To induce the formation of intracytoplasmic membranes (ICM) (greening), cells grown aerobically both in batch culture and in a carbon-limited chemostat were transferred to semiaerobic conditions. In both aerobic cultures, a low level of photosynthetic complexes was observed, which were composed of the reaction center and the LH1 core antenna. Interestingly, in the batch cultures the reaction centers were essentially inactive in forward electron transfer and exhibited low photochemical yields F_V/F_M, whereas the chemostat culture displayed functional reaction centers with a rather rapid (1-2 ms) electron transfer turnover, as well as a high F_V/F_M of ∼0.8. In both cases, the transfer to semiaerobiosis resulted in rapid induction of bacteriochlorophyll a synthesis that was reflected by both an increase in the number of LH1-reaction center and peripheral LH2 antenna complexes. These studies establish that photosynthetic units are assembled in a sequential manner, where the appearance of the LH1-reaction center cores is followed by the activation of functional electron transfer, and finally by the accumulation of the LH2 complexes.     

Photosynthetic oxygen production and the size of the photosynthetic unit in a cell-free preparation from Cyanidium caldarium

A cell-free preparation has been isolated from a mutant of Cyanidium caldarium, grown under conditions such that there is 15 times less chlorophyll per photosynthetic unit than in normal green algae. The preparation is sensitive to 3-(3,4-dichlorophenyl)-1,1-dimethylurea and shows the well-characterized oscillation of O₂ yield, from saturating flashes, following a period of dark adaptation. Greening experiments with dark-grown, wild-type Cyanidium show that the synthesis of photosynthetic units precedes that of bulk chlorophyll and that the O₂-producing system is assembled before the total system coupled to CO₂. No large-scale cooperation of chlorophyll molecules is required for O₂ production.     

The ultrastructure of the marine blue green alga,Trichodesmium erythraeum,with special reference to the cell wall,gas vacuoles,and cylindrical bodies

Summary The marine blue green alga, Trichodesmium erythraeum, was studied with electron microscopy in an attempt to elucidate the structural basis for its rapid lysis when removed from its marine environment. In this connection, it was found that a thining of the electron-dense layer of the longitudinal wall at the site adjacent to transverse wall attachment was responsible for lysis. The underlying biochemical basis for this change has not been elucidated because of the extreme difficulties of maintaining and growing the alga in culture under defined conditions. Several other features of considerable interest also were found. Especially interesting is the very regular array of gas vacuoles in the form of a hollow cylinder which shields most of the photosynthetic system. It was suggested that the gas vacuoles might possibly function optically, having adaptive value in protecting the free-floating alga from excessive radiation. In addition, a detailed structure of the cylindrical bodies was presented, and its structure with the photosynthetic lamellae was compared. On the basis of sectoring to form fragments of double lamellar units from the cylindrical body which are identical in structure to the photosynthetic lamellae, it has been postulated that the cylindrical body may be the site of synthesis for the photosynthetic system in Trichodesmium erythraeum.     

Spectrally decomposed dark-to-light transitions in a PSI-deficient mutant of Synechocystis sp. PCC 6803

Cyanobacterial thylakoid membranes are known to host photosynthetic and respiratory complexes. This hampers a straight forward interpretation of the highly dynamic fluorescence originating from photosynthetic units. The present study focuses on dark-to-light transitions in whole cells of a PSI-deficient mutant of the cyanobacterium Synechocystis sp. PCC 6803. The time-dependent cellular fluorescence spectrum has been measured, while having previously exposed the cells to different conditions that affect respiratory activity. The analysis method used allows the detected signal to be decomposed in a few components that are then assigned to functional emitting species. Additionally, we have worked out a minimal mathematical model consisting of sensible postulated species to interpret the recorded data. We conclude that the following two functional complexes play a major role: a phycobilisome antenna complex coupled to a PSII dimer with either two or no closed reaction centers. Crucially, we present evidence for an additional species capable of strongly quenching fluorescence, whose formation requires the presence of oxygen.     

Structural and functional diversification of the light-harvesting complexes in photosynthetic eukaryotes

Eukaryotes acquired photosynthetic metabolism over a billion years ago, and during that time the light-harvesting antennae have undergone significant structural and functional divergence. The antenna systems are generally used to harvest and transfer excitation energy into the reaction centers to drive photosynthesis, but also have the dual role of energy dissipation. Phycobilisomes formed the first antenna system in oxygenic photoautotrophs, and this soluble protein complex continues to be the dominant antenna in extant cyanobacteria, glaucophytes, and red algae. However, phycobilisomes were lost multiple times during eukaryotic evolution in favor of a thylakoid membrane-integral light-harvesting complex (LHC) antenna system found in the majority of eukaryotic taxa. While photosynthesis spread across different eukaryotic kingdoms via endosymbiosis, the antenna systems underwent extensive modification as photosynthetic groups optimized their light-harvesting capacity and ability to acclimate to changing environmental conditions. This review discusses the different classes of LHCs within photosynthetic eukaryotes and examines LHC diversification in different groups in a structural and functional context.