What does grana do

Chloroplast: An organelle found only in plants and photosynthetic protists that absorbs sunlight and uses it to drive the synthesis of organic compounds from carbon dioxide and water. Granum: plural, grana A stacked portion of the thylakoid membrane in the chloroplast.

Grana function in the light reactions of photosynthesis. Lamella: A sheet like membrane found within a chloroplast of an autotrophic cell. They act as a type of wall at which chloroplasts can be fixed within, achieving the maximum light possible.

Pectin: A carbohydrate consisting of several sugar molecules including rhamnose, galactose, arabinose, and galactouronic acid. Functions in the middle lamella of plant cells to adhere adjacent cells to one another. Functions in the primary cell wall to link adjacent microfibrils.

Pigment: Any material resulting in color in plant or animal cells which is the result of selective absorption. Examples of plant pigments include chlorophyll, xanthophylls and anthocyanins. Plasmodesma: plural, plasmodesmata An open channel in the cell wall of plants through which strands of cytosol connect from adjacent cells.

Plastid: One of a family of closely related plant organelles, including chloroplasts, chromoplasts, and amyloplasts. Stroma: The fluid of the chloroplast surrounding the thylakoid membrane; involved in the synthesis of organic molecules from carbon dioxide and water. Thylakoid: A flattened membrane sac inside the chloroplast, used to convert light energy to chemical energy.

Turgor Pressure: The force directed against a cell wall after the influx of water and the swelling of a walled cell due to osmosis. Autotroph: An organism that obtains organic food molecules without eating other organisms or substances derived from other organisms.

Calvin Cycle: The second of two major stages in photosynthesis, following the light reactions, involving atmospheric CO2 fixation and reduction of the fixed carbon into carbohydrates.

Carbon Fixation: The incorporation of carbon from CO2 into an organic compound by an autotrophic organism. The most abundant protein in prolamellar bodies is the NADPH:protochlorophyllide oxidoreductase POR , and mutants that lack this enzyme accumulate precursors of protochlorophyllide and possess no prolamellar bodies Lebedev et al.

As the initial step in light adaptation, protochlorophyllide is reduced by POR to chlorophyllide and later esterified into chlorophyll Von Wettstein et al. At the same time, the prolamellar body loses its semi-crystalline structure and the extruded lamellae align in parallel throughout the stroma Fig. Whether the lipids of the prolamellar body are directly incorporated into the prothylakoids or are transferred via vesicles is unclear Rosinski and Rosen, ; Adam et al.

The transformation of the semi-crystalline prolamellar body into planar thylakoids Fig. Schematic overview of the light-dependent de-etiolation process and the biogenesis of thylakoid membranes. A The etioplast contains prothylakoids and the semi-crystalline prolamellar body. B Etio-chloroplast stage. The semi-crystalline prolamellar body disassembles.

Whether the lipids of the prolamellar body are incorporated into the maturing prothylakoids directly or via vesicular or tubular intermediates is unknown. Mainly monomeric proteins are incorporated into the developing thylakoids. C The lamellar structures align in parallel within the chloroplast. With the formation of protein complexes, the thylakoids enter a photoactive state.

D The grana stacks characteristic of mature thylakoids form upon incorporation of mega- and supercomplexes. It is assumed that lipids reach thylakoids only via i a vesicular pathway, but might also be supplied by ii soluble glycerolipid transfer proteins or iii via invaginations of, or direct contact sites with, the inner envelope. This schematic representation of the processes involved in the de-etiolation of etioplasts does not do justice to the complexity of the intermediate steps reviewed in Solymosi and Schoefs, For example, grana stacks and prolamellar bodies have been observed in the same chloroplast in electron micrographs Solymosi and Schoefs, The thylakoid membrane contains five major lipids.

Because the synthesis of all of these lipids is finalized in the chloroplast envelope, a mechanism for their continuous transport to thylakoids must exist Jouhet et al. Whether this transfer of lipids occurs via i a vesicular pathway; ii soluble glycerolipid transfer proteins; or iii invaginations that directly connect the envelope to thylakoids is not clear Fig. A mutant with a defective MGDG synthase 1 mgd1 is unable to produce photosynthetically active membranes, but shows invaginations of the inner envelope Kobayashi et al.

Because it is found both at the inner envelope and at the thylakoids Li et al. Other findings hint that VIPP1 might function like its bacterial homologue, namely acting to maintain the chloroplast envelope instead of inducing vesicles, thus having a protective rather than a driving effect on thylakoid biogenesis Zhang et al.

It has been speculated that thylakoid-associated VIPP1 has a similar function, but conclusive proof is still missing Vothknecht et al. Intriguingly, recent results point to a role for VIPP1 in the assembly of thylakoid core complexes Nordhues et al. VIPP1 apparently does not play a role in vesicular transport in mature chloroplasts, but several other proteins remain as candidates for such factors.

Thus, a bioinformatics approach has identified chloroplast-located homologues of the COPII vesicular pathway between the endoplasmic reticulum and Golgi apparatus Andersson and Sandelius, Furthermore, cpSar1 has been detected around cold-induced vesicles Garcia et al. While cpSar1 knock-out mutants show developmental arrest before greening, cpSar1 RNAi RNA interference lines show an interesting intermediate phenotype with respect to thylakoid biogenesis.

In these lines, plastids contain vesicles of various sizes that eventually coalesce and form the typical mature grana stacks Garcia et al. The dynamin family member FZL is also localized at the envelope and thylakoids, and shows GTPase activity, but in FZL knock-out plants disruption of thylakoid ultrastructure is less severe Gao et al.

Although grana stacks are disorganized and vesicles accumulate, FZL is believed to play a more prominent role later in thylakoid development Gao et al. Given the complexity of COPII vesicular transport, a mechanism dedicated solely to the transport of lipids from the envelope to thylakoids is hard to imagine, especially since the non-bilayer-forming nature of MGDG would complicate such mechanisms. Thus, the integration of high concentrations of MGDG into a lipid bilayer relies on the presence of membrane proteins.

However, whether bilayer and non-bilayer phases can co-exist in vesicles too remains speculative. Alternatively, it was suggested that a vesicle pathway might also transport non-lipid components Westphal et al. This idea is in line with the identification of the plasma membrane as the location of initial photosystem biogenesis in cyanobacteria Zak et al.

In an ongoing bioinformatics analysis, the search for components of the vesicular transport mechanism in plastids has been expanded to associated factors Khan et al. In this study, chloroplast-targeted homologues of coat proteins, cargo receptors, tethering factors, and SNAREs were identified.

Despite the lack of conclusive experimental proof, the evidence for a vesicular transport system within the chloroplast cannot be easily dismissed Brandizzi, Thus, it seems likely that thylakoid biogenesis is influenced by the insertion of protein complexes into the lipid bilayer matrix.

Most of the thylakoid proteins are encoded in the nucleus and synthesized in the cytosol, and must be post-translationally imported into the chloroplast. The pathways mediating this transport, and its evolution and regulation, have been extensively reviewed Gutensohn et al.

Recently, it was suggested that lumenal proteins are also essential for thylakoid biogenesis Shipman-Roston et al. Their proper maturation may be a key step in the assembly of thylakoids, as plants mutant for the processing peptidase PLSP1, which is involved in the maturation of lumenal proteins such as OE33, OE23, and plastocyanin , have been shown to accumulate large amounts of vesicles in the stroma but fail to develop intact thylakoids in adult plants Inoue et al.

Here, the critical step seems to be the removal of the thylakoid-transfer signal. Generally, it is difficult to determine unambiguously the importance of integral membrane proteins for thylakoid biogenesis, since their absence results in significant perturbation of photosynthetic activity.

In the following, defects in the assembly of the major thylakoid protein complexes will be reviewed in the context of their effects on thylakoid biogenesis. Mutants without PSI are incapable of photoautotrophic growth. First identified in a series of high chlorophyll fluorescence hcf mutants Meurer et al. Interestingly, other mutants specifically lacking PSI form fragmentary stroma lamellae but still express near wild-type levels of the light-harvesting complexes.

Therefore, it can be concluded that the presence of PSI is essential for thylakoid biogenesis, more specifically the formation of the stroma lamellae. The aforementioned PSII assembly mutant hcf forms enlarged and denser grana stacks, while light-harvesting complexes assemble normally Meurer et al.

Conversely, overexpression of maize plastidial transglutaminase in tobacco increased the numbers of PSII centres in the appressed grana, leading to larger grana stacks and reduced stroma lamellae Ioannidis et al. In atab2 mutants, PSI complexes are absent, PSII is decreased 5-fold, while the Cyt b 6 f and ATPase complexes are expressed normally, leading to a significant decrease in stroma lamellae and a general decrease in thylakoid membrane content Barneche et al.

In the alb3 mutant, a significant loss in thylakoid membrane and grana stacking is observed Sundberg et al. As mentioned above, the impact on grana formation of a specific lack of LHCII trimers or alterations in their subunit composition is less severe. This decrease in grana stacking due to reduced LHCII levels was attributed to a decline in van der Waals attraction, lower electrostatic interaction between opposite charges across the partitioning gap, and impaired formation of PSII—LHCII aggregates, which together appear to exert stronger negative effects on grana formation than the positive effects caused by the weaker electrostatic repulsion due to the lack of LHCII Chow et al.

Mutants affected in the assembly of the Cyt b 6 f complex Lennartz et al. Nevertheless, ultrastructural data are available for a mutant with markedly reduced levels of Cyt b 6 f Manara et al. However, these lines show normal thylakoid formation, indicating that the Cyt b 6 f complex does not play a significant role in the establishment of the thylakoid ultrastructure Manara et al.

The composition of thylakoid membranes varies little between photosynthetically active organisms Siegenthaler, ; Vigh et al. Conversely, proteins can modulate the phase behaviour of MGDG. Thus, by increasing the amounts of LHCII, the inverted hexagonal phase can be progressively transformed into ordered lamellar structures Simidjiev et al.

It has been hypothesized that the amount of thylakoid-incorporated non-bilayer-forming lipid is controlled by the current state of the membrane Garab et al. The existence of such a non-bilayer phase and its exchange with the membrane was shown by Krumova et al. Only the tight packing of proteins into the membrane is compatible with the high concentration of MGDG, and vice versa. We thank Paul Hardy for critical reading of the manuscript.

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