Benning C. A role for lipid trafficking in chloroplast biogenesis. Progress in Lipid Research 47 , — Mechanisms of lipid transport involved in organelle biogenesis in plant cells. Annual Review of Cell and Developmental Biology 25 , 71 — Molecular Plant 2 , — Light-induced dissociation of an antenna hetero-oligomer is needed for non-photochemical quenching induction.
Proteomic analysis of highly purified prolamellar bodies reveals their significance in chloroplast development. Photosynthesis Research 96 , 37 — Photosystem II core phosphorylation and photosynthetic acclimation require two different protein kinases.
Glycerolipids in photosynthesis: composition, synthesis and trafficking. Brandizzi F. Traffic 12 , 9 — A note on three-dimensional models of higher-plant thylakoid networks. The Plant Cell 20 , — ; author reply — Identification of a functional respiratory complex in chloroplasts through analysis of tobacco mutants containing disrupted plastid ndh genes.
EMBO Journal 17 , — C-terminal processing of reaction center protein D1 is essential for the function and assembly of photosystem II in Arabidopsis.
Surface charges, the heterogeneous lateral distribution of the two photosystems, and thylakoid stacking. Biochimica et Biophysica Acta , 69 — Granal stacking of thylakoid membranes in higher plant chloroplasts: the physicochemical forces at work and the functional consequences that ensue. Photochemical and Photobiological Sciences 4 , — The Plant Cell 20 , — Cell , — Electron tomography of plant thylakoid membranes.
Journal of Experimental Botany 62 , — The Plant Cell 22 , — EMBO Journal 22 , — Investigations of the role of the main light-harvesting chlorophyll—protein complex in thylakoid membranes. Reconstitution of depleted membranes from intermittent-light-grown plants with the isolated complex.
Journal of Cell Biology 98 , — Supramolecular organization of thylakoid membrane proteins in green plants. Biochimica et Biophysica Acta , 12 — Changes in light energy distribution upon state transitions: an in vivo photoacoustic study of the wild type and photosynthesis mutants from Chlamydomonas reinhardtii.
Photosystem II supercomplex remodeling serves as an entry mechanism for state transitions in Arabidopsis. The Plant Cell 23 , — Lateral diffusion of an integral membrane protein: Monte Carlo analysis of the migration of phosphorylated light-harvesting complex II in the thylakoid membrane.
Biochemistry 32 , — Functional analysis of a divergent system II protein, Ccs1, involved in c-type cytochrome biogenesis. PhosPhAt: the Arabidopsis thaliana phosphorylation site database. An update. Nucleic Acids Research 38 , D — D Chloroplast-targeted Hsp90 plays essential roles in plastid development and embryogenesis in Arabidopsis possibly linking with VIPP1.
Physiologia Plantarum , — Franklin KA. Shade avoidance. New Phytologist , — Prerequisites for terminal processing of thylakoidal Tat substrates. The Plant Cell 21 , — Self-regulation of the lipid content of membranes by non-bilayer lipids: a hypothesis. Trends in Plant Science 5 , — Alterations in chloroplast thylakoids during an in vitro freeze—thaw cycle. Plant Physiology 57 , — The chloroplast protein CPSAR1, dually localized in the stroma and the inner envelope membrane, is involved in thylakoid biogenesis.
The Plant Journal 63 , 73 — Visualizing the mobility and distribution of chlorophyll proteins in higher plant thylakoid membranes: effects of photoinhibition and protein phosphorylation. The Plant Journal 62 , — Light-harvesting antenna composition controls the macrostructure and dynamics of thylakoid membranes in Arabidopsis.
The Plant Journal 69 , — Goss R Wilhelm C. Lipids in algae, lichens and mosses. In: Wada H Murata N , eds. Lipids in photosynthesis: essential and regulatory functions , Vol.
Dordrecht, The Netherlands : Springer , — The thylakoid membranes of higher plant chloroplasts. Biochemical Journal , — Lateral heterogeneity of polar lipids in the thylakoid membranes of spinach chloroplasts. Toc, Tic, Tat et al. Journal of Plant Physiology , — The PSI-F subunit is essential for photoautotrophic growth and contributes to antenna function. Architectural switch in plant photosynthetic membranes induced by light stress. PGRL1 is the elusive ferredoxin-plastoquinone reductase in photosynthetic cyclic electron flow.
Molecular Cell 49 , — Lipid traffic: floppy drives and a superhighway. Nature Reviews Molecular Cell Biology 6 , — Journal of Molecular Biology , — Horton P.
Are grana necessary for regulation of light harvesting? Molecular Plant 6 , — Complete maturation of the plastid protein translocation channel requires a type I signal peptidase.
Journal of Cell Biology , — Leaf positioning of Arabidopsis in response to blue light. Molecular Plant 1 , 15 — Remodeling of tobacco thylakoids by over-expression of maize plastidial transglutaminase. Izawa S Good NE. Effect of salts and electron transport on the conformation of isolated chloroplasts. Electron microscopy. Plant Physiology 41 , — Understanding the roles of the thylakoid lumen in photosynthesis regulation.
Frontiers in Plant Science 4 , Photoprotective energy dissipation involves the reorganization of photosystem II light-harvesting complexes in the grana membranes of spinach chloroplasts. Glycerolipid transfer for the building of membranes in plant cells.
Progress in Lipid Research 46 , 37 — Kato Y Sakamoto W. Protein quality control in chloroplasts: a current model of D1 protein degradation in the photosystem II repair cycle. Journal of Biochemistry , — Psb29, a conserved kD protein, functions in the biogenesis of Photosystem II complexes in Synechocystis and Arabidopsis.
The Plant Cell 17 , — New putative chloroplast vesicle transport components and cargo proteins revealed using a bioinformatics approach: an Arabidopsis model. PLoS One 8 , e Quality control of photosystem II: thylakoid unstacking is necessary to avoid further damage to the D1 protein and to facilitate D1 degradation under light stress in spinach thylakoids.
Kirchhoff H. Photosynthesis Research , — Structural constraints for protein repair in plant photosynthetic membranes. Plant Signaling and Behavior 8 , e Diffusion of molecules and macromolecules in thylakoid membranes. Low-light-induced formation of semicrystalline photosystem II arrays in higher plant chloroplasts. Biochemistry 46 , — Dynamic control of protein diffusion within the granal thylakoid lumen.
Molecular architecture of the thylakoid membrane: lipid diffusion space for plastoquinone. Biochemistry 41 , — Plastid signalling to the nucleus: messengers still lost in the mists? Trends in Genetics 25 , — Role of galactolipid biosynthesis in coordinated development of photosynthetic complexes and thylakoid membranes during chloroplast biogenesis in Arabidopsis. The Plant Journal 73 , — Supramolecular organization of photosystem II in green plants.
Biochimica et Biophysica Acta , 2 — VIPP1, a nuclear gene of Arabidopsis thaliana essential for thylakoid membrane formation. Phase behavior of phosphatidylglycerol in spinach thylakoid membranes as revealed by 31P-NMR. Lateral mobility of the light-harvesting complex in chloroplast membranes controls excitation-energy distribution in higher-plants. Archives of Biochemistry and Biophysics , — Constitutive expression of pea Lhcb 1—2 in tobacco affects plant development, morphology and photosynthetic capacity.
Plant Molecular Biology 55 , — The Plant Cell 7 , — Lee AG. Current Biology 10 , R — R HCF encodes a thioredoxin-like protein involved in the biogenesis of the cytochrome b 6 f complex in Arabidopsis. The Plant Cell 13 , — Planta , — The universally conserved HCF protein is involved in assembly of [4Fe—4S]-cluster-containing complexes in Arabidopsis thaliana chloroplasts.
The Plant Journal 37 , — Molecular cloning of a chloroplastic protein associated with both the envelope and thylakoid membranes. Plant Molecular Biology 25 , — The Plant Journal 50 , — Molecular Biology of the Cell 16 , — PsbP-domain protein1, a nuclear-encoded thylakoid lumenal protein, is essential for photosystem I assembly in Arabidopsis. The Plant Cell 24 , — Role of vesicle-inducing protein in plastids 1 in cpTat transport at the thylakoid.
The Plant Journal 71 , — Knock-out of the genes coding for the Rieske protein and the ATP-synthase delta-subunit of Arabidopsis. Effects on photosynthesis, thylakoid protein composition, and nuclear chloroplast gene expression.
Isolation of high-chlorophyll-fluorescence mutants of Arabidopsis thaliana and their characterisation by spectroscopy, immunoblotting and northern hybridisation. A nuclear-encoded protein of prokaryotic origin is essential for the stability of photosystem II in Arabidopsis thaliana. Midorikawa T Inoue K. All rights reserved. Part B: courtesy of M. Feist, University of Montpellier. Coleochaete orbicularis. Both the gametophyte and the background are bright green.
The gametophyte has an irregular circular shape and a scalloped edge. It is divided into many box-like segments cells , each with a visible, round nucleus inside.
Panel b shows a Chara gametophyte. The organism has branching, tendril-like leaves reaching from a primary stalk. The green leaves are punctuated with small, round, yellow structures. A green liverwort gametophyte, In panel c, is protruding from the soil.
Its four primary stems each diverge into two halves and then branch again at their termini, so that each has a forked end. Panel d shows a hornwort gametophyte. Each green stem resembles a single blade of grass. Panel e shows moss gametophytes with sporophytes protruding from the ground. The gametophytes have small green leaves, and the sporophytes are thin, unbranched, brown stalks. Each sporophyte has a fluorescent orange, oviform capsule called a sporangia perched on top of its stalk.
Panel f shows six clubmoss sporophytes emanating from the ground. Some stand vertically out of the soil, and some curve or have fallen horizontally. They have many stiff, protruding, spine-like, green leaves. The sporangia are small yellow balls at the base of the leaves.
Panel g shows fern sporophytes with many stems covered with small, elongated, symmetrical green leaves. Panel h shows a whisk fern sporophyte with long, straight, green stems beaded with yellow, round synangia along their lengths. In panel i, a horsetail sporophyte is shown. It has a single long stem, which is surrounded by a skirt of green leaves at its base and an elongated, yellow cone at the top.
In Panel j, a large Cycas seed plant sporophyte is shown. Long fronds emanate upwards from the plant's trunk, and in the center of them there is a large mass called the cone.
Panel a is a photomicrograph of a gametophyte of a microscopic green alga called Coleochaete orbicularis. Most living things depend on photosynthetic cells to manufacture the complex organic molecules they require as a source of energy. Photosynthetic cells are quite diverse and include cells found in green plants, phytoplankton, and cyanobacteria. During the process of photosynthesis, cells use carbon dioxide and energy from the Sun to make sugar molecules and oxygen.
These sugar molecules are the basis for more complex molecules made by the photosynthetic cell, such as glucose. Then, via respiration processes, cells use oxygen and glucose to synthesize energy-rich carrier molecules, such as ATP, and carbon dioxide is produced as a waste product.
Therefore, the synthesis of glucose and its breakdown by cells are opposing processes. Figure 2 2 in the sky represents the process of photosynthesis. Two arrows are directed outwards from the trees towards the atmosphere. One represents the production of biomass in the trees, and the other represents the production of atmospheric carbon dioxide CO 2.
Arrows emanating from a tree's roots point to two molecular structures: inorganic carbon and organic carbon, which may decompose into inorganic carbon. The space enclosed by the inner chloroplast membrane is called the stroma. Chloroplasts replicate giving rise to new chloroplasts as they grow and divide. They also have their own DNA and ribosomes. The thylakoid membranes contain the pigments chlorophyll and carotenoids, as well as enzymes and the electron transport chains used in photosynthesis def , a process that converts light energy into the chemical bond energy of carbohydrates.
Energy trapped from sunlight by chlorophyll is used to excite electrons in order to produce ATP by photophosphorylation. The light-independent reactions of photosynthesis use this ATP and NADPH to produce carbohydrates from carbon dioxide and water, a series of reactions that occur in the stroma of the chloroplast.
Study the material in this section and then write out the answers to these questions. Do not just click on the answers and write them out. It is not known however, why different plant species have different arrangements of grana within their chloroplasts. In pea, larger appressed thylakoids are regularly arranged within chloroplasts as uniformly distributed red fluorescent bodies, while irregular appressed thylakoid membranes within bean chloroplasts correspond to smaller and less distinguished fluorescent areas in CLSM images.
Structural differences influenced the PSII photochemistry, however without significant changes in photosynthetic efficiency. Qualitative and quantitative analysis of chlorophyll-protein complexes as well as spectroscopic investigations indicated a similar proportion between PSI and PSII core complexes in pea and bean thylakoids, but higher abundance of LHCII antenna in pea ones.
Based on proteomic and spectroscopic investigations we postulate that the differences in the chloroplast structure between the analyzed species are a consequence of quantitative proportions between the individual CP complexes and its arrangement inside membranes.
Such a structure of membranes induced the formation of large stacked domains in pea, or smaller heterogeneous regions in bean thylakoids. Presented 3D models of chloroplasts showed that stacked areas are noticeably irregular with variable thickness, merging with each other and not always parallel to each other. The thylakoid system in plants is organized into two distinct domains: grana arranged in stacks of appressed membranes and non-appressed membranes consisting of stroma thylakoids and margins of granal stacks [ 1 ].
It is known that appressed membranes that form grana are not essential for photosynthesis but they are ubiquitous in all chlorophyll Chl b -containing higher plants [ 1 , 2 ].
Many photosynthetic organisms such as red algae, Cyanobacteria, many green algae such as Chlamydomonas reinhardtii have no stacked thylakoids. Apart from higher plants only Charophyta have appressed membranes indistinguishable from those of land plants [ 1 ]. Why did plants develop grana? The development of appressed membranes caused structural heterogeneity that is reflected by functional differentiation with respect to the location of hierarchically organized photosyntetic complexes in supercomplexes and megacomplexes within appressed and non-appressed membranes [ 3 , 4 ].
The structural and organizational changes of grana stacks are driven by physical and chemical forces. It is believed that membrane appression is maintained primarily by the balance between the van der Waals attraction versus electrostatic and hydratation repulsion [ 6 ].
It is argued that the reason for the development of appressed membranes in plants is that their photosynthetic apparatus needs to cope with and survive ever-changing environmental conditions, such as transition from darkness, low-light to high-light conditions [ 1 ] or temperature fluctuation [ 11 , 12 ]. Short-term changes are due to the redistribution of absorbed excitation energy state to state transition that is based on migration of LHCII from PSII after its phosphorylation [ 13 — 15 ].
Why do different plant species have different arrangements of grana within their chloroplasts? Examinations of mutants yielded some information on the arrangement of thylakoid membranes within chloroplasts. For example it is known that Arabidopsis Aba mutants — deficient in epoxy-carotenoids — have significantly more grana stacks per chloroplast and more chloroplasts per cell but reduced thylakoid stacking in comparison with wild Arabidopsis plants [ 16 ].
In tobacco knockdown of PsbP protein, one of the three oxygen evolving complex proteins in plants, impairs the accumulation of PSII supercomplexes in tobacco and causes large disorder in the thylakoid grana stacking [ 17 ]. Studies of the arrangements of thylakoid membranes gave information on chlorophyll b —less Arabidopsis mutants. Markedly decreased level or the absence of most of Lhcbs caused fewer grana and much longer stromal thylakoids than in the wild type which implied that the total granal cross-sectional area per chloroplast area was decreased dramatically [ 18 ].
Ruban and coworkers [ 19 ] argued that in the absence of one of the PSII supercomplex main components, another antenna protein may be recruited to replace a mutated or absent protein, therefore allowing the main complex to assemble and function correctly. Mutations in the photosynthetic apparatus that cause changes in the thylakoid arrangement give an altered picture of chloroplast thylakoid membranes with profound effect on the photosynthetic efficiency and capacity [ 18 ].
Data on the thylakoid arrangement in chloroplasts of different plant species grown in natural or varying environmental conditions, can supplement information obtained for the mutants. In this paper we propose possible interpretation at molecular level of different arrangements of the thylakoid membranes of pea Pisum sativum L. Here we compared different chloroplast structures by confocal laser scanning microscopy CLSM followed by computer modeling and by transmission electron microscopy TEM.
By applying these diverse methodologies we attempted to describe the relationship between spatial chloroplast structure detected by CLSM in situ and the arrangements of CP complexes within the thylakoid membranes. CLSM images revealed fluorescent red spots of about 0.
Chlorophyll fluorescence of intact chloroplasts revealed by CLSM and 3D reconstruction of chloroplast structure.
Each red image presents the maximum intensity projection of deconvolved stack of CLSM images. Images are representative for at least 20 independent experiments. Grey images represent 3D models of intact pea C, E and bean chloroplasts D, F created after deconvolution.
Each image is a representative of at least 10 independent experiments. Large number 94— of fluorescence images were taken in different focal depths. The surface of Chl fluorescence areas is seen in the face view as well as in the side view.
For a better view of chloroplast 3D structures, animated models are included in Additional file 1 : Video S1 and Additional file 2 : Video S2. The 3D structures reflect the distribution of Chl fluorescence in a chloroplast as a whole.
These appressed regions in TEM images of bean chloroplasts are counterparts of smaller and less distinguished fluorescent areas seen by CLSM cf. Mesophyll chloroplast images revealed by TEM. Pictures show different thylakoid arrangements in pea A and bean chloroplasts D , larger appressed thylakoids regions in pea B than in bean E and wider thylakoid lumen in pea C than in bean F.
The average value of the ratio of the length of grana thylakoids to the length of stroma thylakoids was 3. The distance between the pairs of membranes in thylakoids was estimated to be The chloroplast images from intact chloroplasts by CLSM and from leaf tissue in situ by TEM were consistent with each other and gave similar information regarding different distribution and appearance of appressed and non-appressed thylakoids in pea and bean chloroplasts.
More than 50 proteins from 10 to kD in molecular weight were distinguishable Additional file 3 : Figure S1. Protein analysis of pea and bean thylakoid membranes. Separated bands were analyzed by immunodetection and identified by mass spectrometry. Molecular weights [kD] of pea and bean Lhca and Lhcb are given in the table C.
However significant compositional differences were observed in the gel area where the antennae proteins Lhca and Lhcb were localized 20—30 kD. Quantitative relationships between the two samples were expressed as a ratio of pixel intensities corresponding to selected proteins bands normalized to chlorophyll content.
Lhcb1 was detected as one major band in pea and one major and two minor bands in bean samples. The presence of these two additional bands in bean sample is probably an effect of anti- Lhcb1 antibody cross interactivity with the Lhcb4 top band and the Lhcb3 bottom band proteins cf. On the other hand the level of PsbA in bean was about two times higher than in pea, which suggests a higher ratio of Lhcb1 to PsbA in pea thylakoids.
The Lhca1 detection pattern consisted of one dominant and three pea or six bean minor bands with higher molecular weight. PsaA, the PSI core protein, was detected as 55—60 kD band whereas expected molecular weight for this protein is around 83 kD [ 24 ]. The PsaA level in pea thylakoids is similar to bean up to the standard deviation. Pea and bean thylakoid samples of increasing amount of chlorophyll starting from 0.
Due to statistically significant difference in Lhcb1 and PsbA ratios we can explicitly conclude that the proportion of PSII antennae to the core proteins was higher in pea than in bean thylakoids. The pea and bean complexes differed in the electrophoretic mobility of their components, especially of CP7 to CP10 that appeared to have higher molecular weights in pea thylakoids.
Immunodetection of proteins that are the part of CP bands were resolved by mild-denaturing electrophoresis B. In the monomer region CP8-CP10 many of individual proteins were resolved. Quantitative analysis of mild-denaturing electrophoresis pattern showed significant differences in chlorophyll-protein complexes isolated from pea and bean leaves. Because the mild-denaturing electrophoresis did not preserve completely the CP supercomplexes structure [ 25 ], the presented results illustrate the stability of distinct CP complexes after the detergent treatment rather than its native organization in thylakoid membranes.
The CP core complexes bind Chl a only [ 4 , 8 ]. The thylakoid emission spectrum is highly heterogeneous and might be resolved in some partially overlapping emission bands corresponding to specific CP complexes [ 10 , 28 ]. The best Gaussian fit to fluorescence spectra enable to distinguish five components Additional file 4 : Figure S2. The spectra A, B were normalized to the area of under the spectrum, and the difference spectra C, D were calculated for the respective excitation spectra.
The presented spectra are representative of three separate experiments. These data indicate distinct differences in the LHCI-PSI structure between the two species; probably due to the presence of a number of Lhca isoforms cf. Steady-state electron absorption spectra revealed the properties of pigments bound in CP complexes [ 33 ].
Normalized excitation spectra presented the relative energy transfer from the absorbing pigments to the emitting Chl species reflecting the state of CP complexes [ 33 ].
0コメント