thylakoid pattern
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While photosynthetic processes have become increasingly understood in cyanobacterial model strains, differences in the spatial distribution of thylakoid membranes among various lineages have been largely unexplored. Cyanobacterial cells exhibit an intriguing diversity in thylakoid arrangements, ranging from simple parietal to radial, coiled, parallel, and special types. Although metabolic background of their variability remains unknown, it has been suggested that thylakoid patterns are stable in certain phylogenetic clades. For decades, thylakoid arrangements have been used in cyanobacterial classification as one of the crucial characters for definition of taxa. The last comprehensive study addressing their evolutionary history in cyanobacteria was published 15 years ago. Since then both DNA sequence and electron microscopy data have grown rapidly. In the current study, we map ultrastructural data of >200 strains onto the SSU rRNA gene tree, and the resulting phylogeny is compared to a phylogenomic tree. Changes in thylakoid architecture in general follow the phylogeny of housekeeping loci. Parietal arrangement is resolved as the original thylakoid organization, evolving into complex arrangement in the most derived group of heterocytous cyanobacteria. Cyanobacteria occupying intermediate phylogenetic positions (greater filamentous, coccoid, and baeocytous types) exhibit fascicular, radial, and parallel arrangements, partly tracing the reconstructed course of phylogenetic branching. Contrary to previous studies, taxonomic value of thylakoid morphology seems very limited. Only special cases such as thylakoid absence or the parallel arrangement could be used as taxonomically informative apomorphies. The phylogenetic trees provide evidence of both paraphyly and reversion from more derived architectures in the simple parietal thylakoid pattern. Repeated convergent evolution is suggested for the radial and fascicular architectures. Moreover, thylakoid arrangement is constrained by cell size, excluding the occurrence of complex architectures in cyanobacteria smaller than 2 μm in width. It may further be dependent on unknown (eco)physiological factors as suggested by recurrence of the radial type in unrelated but morphologically similar cyanobacteria, and occurrence of special features throughout the phylogeny. No straightforward phylogenetic congruences have been found between proteins involved in photosynthesis and thylakoid formation, and the thylakoid patterns. Remarkably, several postulated thylakoid biogenesis factors are partly or completely missing in cyanobacteria, challenging their proposed essential roles.
- Klíčová slova
- SSU rRNA gene, cyanobacteria, evolution, photosynthesis, phylogenomics, taxonomy, thylakoid pattern,
- Publikační typ
- časopisecké články MeSH
Since the publication of the fluid-mosaic membrane theory by Singer and Nicolson in 1972 generations of scientists have adopted this fascinating concept for all biological membranes. Assuming the membrane as a fluid implies that the components embedded in the lipid bilayer can freely diffuse like swimmers in a water body. During the detailed biochemical analysis of the thylakoid protein components of chloroplasts from higher plants and algae, in the '80 s and '90 s it became clear that photosynthetic membranes are not homogeneous either in the vertical or the lateral directions. The lateral heterogeneity became obvious by the differentiation of grana and stroma thylakoids, but also the margins have been identified with a highly specific protein pattern. Further refinement of the fluid mosaic model was needed to take into account the presence of non-bilayer lipids, which are the most abundant lipids in all energy-converting membranes, and the polymorphism of lipid phases, which has also been documented in thylakoid membranes. These observations lead to the question, how mobile the components are in the lipid phase and how this ordering is made and maintained and how these features might be correlated with the non-bilayer propensity of the membrane lipids. Assuming instead of free diffusion, a "controlled neighborhood" replaced the model of fluidity by the model of a "mixed crystal structure". In this review we describe why basic photosynthetic regulation mechanisms depend on arrays of crystal-like lipid-protein macro-assemblies. The mechanisms which define the ordering in macrodomains are still not completely clear, but some recent experiments give an idea how this fascinating order is produced, adopted and maintained. We use the operation of the xanthophyll cycle as a rather well understood model challenging and complementing the standard Singer-Nicolson model via assigning special roles to non-bilayer lipids and non-lamellar lipid phases in the structure and function of thylakoid membranes.
- Klíčová slova
- Fluid mosaic model, Lipid bilayer, Membrane model, Photosynthesis, Thylakoid,
- MeSH
- difuze MeSH
- fotosyntéza * MeSH
- intracelulární membrány chemie metabolismus MeSH
- membránové lipidy metabolismus MeSH
- tylakoidy chemie metabolismus MeSH
- Publikační typ
- časopisecké články MeSH
- přehledy MeSH
- Názvy látek
- membránové lipidy MeSH
Non-photochemical quenching (NPQ) is an important photoprotective mechanism in plants and algae. Although the process is extensively studied, little is known about its relationship with ultrastructural changes of the thylakoid membranes. In order to better understand this relationship, we studied the effects of illumination on the organization of thylakoid membranes in Monstera deliciosa leaves. This evergreen species is known to exhibit very large NPQ and to possess giant grana with dozens of stacked thylakoids. It is thus ideally suited for small-angle neutron scattering measurements (SANS)-a non-invasive technique, which is capable of providing spatially and statistically averaged information on the periodicity of the thylakoid membranes and their rapid reorganizations in vivo. We show that NPQ-inducing illumination causes a strong decrease in the periodic order of granum thylakoid membranes. Development of NPQ and light-induced ultrastructural changes, as well as the relaxation processes, follow similar kinetic patterns. Surprisingly, whereas NPQ is suppressed by diuron, it impedes only the relaxation of the structural changes and not its formation, suggesting that structural changes do not cause but enable NPQ. We also demonstrate that the diminishment of SANS peak does not originate from light-induced redistribution and reorientation of chloroplasts inside the cells.
- Klíčová slova
- chloroplast thylakoid membranes, grana, lamellar repeat distance, non-photochemical quenching, small-angle neutron scattering,
- MeSH
- Araceae chemie MeSH
- difrakce rentgenového záření * MeSH
- intracelulární membrány chemie metabolismus MeSH
- listy rostlin chemie MeSH
- maloúhlový rozptyl * MeSH
- neutrony * MeSH
- světlo MeSH
- tylakoidy genetika metabolismus MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Isolated thylakoids from halophytic Eutrema salsugineum (Thellungiella salsuginea) produces more H2O2 in comparison to glycophytic Arabidopsis thaliana. The first objective of this study was to verify whether this feature is relevant also to the intact chloroplasts and leaves. Enhanced H2O2 levels in chloroplasts and leaves of E. salsugineum were positively verified with several methods (electron microscopy, staining with Amplex Red and with diaminobenzidine). This effect was associated with a decreased ratio of [Formula: see text]/H2O2 in E. salsugineum in comparison to A. thaliana as detected by electron paramagnetic resonance method. As a next step, we tested how this specific ROS signature of halophytic species affects the antioxidant status and down-stream components of ROS signaling. Comparison of enzymatic antioxidants revealed a decreased activity of ascorbate peroxidase (APX), enhanced activity of glutathione peroxidase, and the presence of thylakoid-bound forms of iron superoxide dismutase (FeSOD) and APX in E. salsugineum. These cues were, however, independent from application of salt stress. The typical H2O2-dependent cellular responses, namely the levels of glucosinolates and stress-related hormones were determined. The total glucosinolate content in E. salsugineum water-treated leaves was higher than in A. thaliana and increased after salinity treatment. Treatment with salinity up-regulated all of tested stress hormones, their precursors and catabolites [abscisic acid (ABA), dihydrophaseic acid, phaseic acid, 1-aminocyclopropane-1-carboxylic acid, salicylic acid, jasmonic acid, cis-(+)-12-oxo-phytodienoic acid and jasmonoyl-L-isoleucine] in A. thaliana, whereas in E. salsugineum only a stimulation in ethylene synthesis and ABA catabolism was noted. Obtained results suggest that constitutively enhanced H2O2 generation in chloroplasts of E. salsugineum might be a crucial component of stress-prepardeness of this halophytic species. It shapes a very efficient antioxidant protection (in which glucosinolates might play a specific role) and a fine tuning of hormonal signaling to suppress the cell death program directed by jasmonate pathway.
- Klíčová slova
- chloroplast, glucosinolates, halophyte, hydrogen peroxide, salinity, stress hormones,
- Publikační typ
- časopisecké články MeSH
Cells of the psbH deletion mutant IC7 of the cyanobacterium Synechocystis PCC 6803 grown in the absence of glucose contain strongly reduced levels of chlorophyll when compared with cells grown in the presence of glucose, or compared with wild-type (WT) cells. Low-temperature fluorescence emission spectra revealed decreased content of both active PS II (Photosystem II) and PS I (Photosystem I) complexes. Analysis of thylakoid membrane complexes of IC7 by native electrophoresis showed a similar set of chlorophyll-proteins, namely a PS II core complex and trimeric and monomeric PS II complexes, as in WT. However, in contrast to WT, the (35)S-methionine protein labeling pattern of the mutant exhibited no preferential labeling of the D1 protein in the PS II core complexes, and the labeled D1 and D2 proteins accumulated predominantly in the PS II reaction center lacking CP47. The results show that in autotrophically grown cells of the psbH deletion mutant, selective D1 turnover is inhibited and synthesis of CP47 becomes a limiting step in the PS II assembly.
- MeSH
- delece genu * MeSH
- fosfoproteiny nedostatek genetika metabolismus MeSH
- fotosystém II (proteinový komplex) biosyntéza chemie genetika metabolismus MeSH
- glukosa metabolismus MeSH
- světlosběrné proteinové komplexy biosyntéza MeSH
- Synechocystis cytologie genetika metabolismus MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- Názvy látek
- fosfoproteiny MeSH
- fotosystém II (proteinový komplex) MeSH
- glukosa MeSH
- photosystem II, chlorophyll-binding protein, CP-47 MeSH Prohlížeč
- photosystem II, psbH subunit MeSH Prohlížeč
- světlosběrné proteinové komplexy MeSH
The in vivo substitution of Mg2+ in chlorophyll by heavy metals is an important damage mechanism in heavy metal-stressed plants that leads to an inhibition of photosynthesis. In photosynthetic organisms with LHC II antennae, the in vivo substitution of Mg2+ by Cu2+ occurs particularly readily under low irradiance with a dark phase - a phenomenon referred to as 'shade reaction'. In the present study the limiting steps of the shade reaction were investigated with synchronised cultures of the chlorococcal green alga Scenedesmus quadricauda (Turp.) Bréb. The rate of copper chlorophyll formation during shade reaction was shown to be controlled by several factors; firstly, in some phases of the cell cycle, especially at the end of the light period, Mg2+ in chlorophyll was not accessible to substitution. This pattern is likely to be caused by cell cycle-dependent changes in photosynthesis and thylakoid ultrastructure, which were published earlier and are reconsidered in the discussion of the present work. Secondly, prolonged culture in a medium containing 3 μM Cu2+ reversibly increased the resistance of the strain to Cu2+. Culturing without added Cu2+ lowered the threshold concentrations of various deleterious effects more than 10-fold within 8 months of de-adaptation. Adaptation to high Cu2+ levels is discussed in the context of studies of the regulation of metal transporter proteins. In addition, it was also observed that toxic Cu2+ levels impaired photosynthesis sooner than cell division.
- Publikační typ
- časopisecké články MeSH
Foliar anthocyanins shape a peculiar shade in a red leaf's interior leading to uneven energy distribution between the two photosystems. Accordingly, a readjustment of PSII/PSI stoichiometry could restore excitation balance. To test this hypothesis, 77 K fluorescence emission spectra of thylakoids from green and red leaves of seven species with different pigment profiles were compared. The ratio of F686/F736 served as an indication of the PSII/PSI functional ratio. To avoid possible species-dependent differences in the measured parameters, plants showing intra-individual, intra-species, or intra-leaf variation in the expression of the anthocyanic character were used. Red leaves or red leaf areas displayed higher PSII/PSI ratio, irrespectively of species and anthocyanin accumulation pattern. PSII/PSI ratio declined in parallel with anthocyanin decrease. In five species, red leaves displayed also a lower Chl a/b ratio. We conclude that red leaves growing in full sunlight develop adaptive adjustments in their chlorophyll and photosystem ratios, compatible with the shade-acclimation syndrome.
- Klíčová slova
- fluorescence emission spectra, foliar anthocyanins, photosystem ratio, shade-acclimation syndrome,
- Publikační typ
- časopisecké články MeSH
The function of selenium in an organism is mediated mostly by selenoproteins including glutathione peroxidase. Glutathione peroxidase is a potent anti-oxidative enzyme, scavenging a variety of peroxides. The green alga Scenedesmus quadricauda was used to investigate the relationship between the toxicity of selenium and the glutathione peroxidase activity. Selenium resistant strains SeIV and SeVI were synchronized and grown in high concentrations of Se (selenite or selenate). As a measure of selenium toxicity the EC(50) values were determined. During growth of the untreated wild type, glutathione peroxidase activity increased slightly and then declined gradually until the end of the cell cycle. A similar pattern was observed in untreated resistant strains and when resistant strains were grown in the presence of selenium in the oxidation state to which they were resistant. In the wild type cultivated with 50 mg Se L(-1) (selenite or selenate), activity increased to a high level and slowly declined until the end of the cell cycle. Similarly, activity increased in strains SeIV and SeVI when grown in the presence of selenium in the oxidation state to which they were not resistant. We followed the effect of selenium on the ultrastructure of S. quadricauda. After exposure to selenite, the chloroplast membranes of wild type were reorganized into thick bundles of thylakoids and the stroma became granulose. When selenate was added, the chloroplast of wild type had a fingerprint-like appearance, the stroma became less dense and starch production increased. In selenium resistant strains, when treated with the selenium form to which they were resistant, the chloroplast was affected, but not to such an extent as in the wild type. The activity of glutathione peroxidase in Scenedesmus was affected by selenium in an oxidation state-dependent manner. The most apparent effects of selenium on the ultrastructure involved impairment of the chloroplast and the overproduction of starch.
- MeSH
- buněčný cyklus účinky léků fyziologie MeSH
- chloroplasty účinky léků metabolismus ultrastruktura MeSH
- fyziologický stres MeSH
- geneticky modifikované rostliny MeSH
- glutathionperoxidasa metabolismus MeSH
- kultivační techniky MeSH
- kyselina selenová MeSH
- Scenedesmus cytologie účinky léků enzymologie fyziologie MeSH
- selen aplikace a dávkování analýza toxicita MeSH
- seleničitan sodný aplikace a dávkování toxicita MeSH
- sloučeniny selenu aplikace a dávkování toxicita MeSH
- testy toxicity MeSH
- vztah mezi dávkou a účinkem léčiva MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- Názvy látek
- glutathionperoxidasa MeSH
- kyselina selenová MeSH
- selen MeSH
- seleničitan sodný MeSH
- sloučeniny selenu MeSH