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See paper for distribution of PS-I and PS-II within grana

As explained in the above paper, the PS-I complex is primarily located within non-appressed thylakoid region and stroma lamella, while PS-II in the appressed regions. Now since non-cyclic electron transfer (non-cyclic photophosphorylation) requires both the photosystems, wouldn't their distribution within grana far from one another hamper their ability to carry out non-cyclic electron transport. Although there are mobile untis to account for the electron transport between marginal PS-I and the PS-II immediately below, but the electron transport to the PS-II units deep in the granal core would still be unaccounted for.

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    $\begingroup$ You may find useful this article. $\endgroup$
    – Mattia Rovetta
    Commented Jun 19, 2013 at 17:59
  • $\begingroup$ I commented instead of answer because I hadn't understood your question precisely! Maybe the problem is mine, but can you refine this phrase?: "Although there are mobile carriers, but what about the PS-II units far in the centre of large stacks of thylakoids?" $\endgroup$
    – Mattia Rovetta
    Commented Jun 23, 2013 at 9:10
  • $\begingroup$ "Although there are mobile untis to account for the electron transport between marginal PS-I and the PS-II immediately below, but the electron transport to the PS-II units deep in the granal core would still be unaccounted for." Edited in answer too. $\endgroup$
    – stochastic13
    Commented Jun 23, 2013 at 9:28

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First a clarification: the mobile units are no so "mobile". Due to (1) the high protein:lipid ratio of the thylakoid membrane and (2) the packing of protein complexes, the rapid diffusion of components along the membrane is hindered.

It has been proposed the existence of small local PQ diffusion microdomains, based upon the percolation theory:

From the percolation theory, above a critical density, immobile or slowly moving transmembrane proteins are, indeed, expected to form flexible 2D networks of closed compartments preventing long range movement of small molecules. (Kirchhoff, H et al. (2000) Biochim. Biophys. Acta 1459, 148–168)

For this reason the PQ are confined, on a short time scale, within these domains, preventing their "dispersion".

In addition of the low diffusion of mobile carriers, the geometry of the granal stacks mantains the contact area between the PSII of the appressed grana and the PSI in the margin annulus domains relatively large, enabling effective linear electron transport in microdomains.

Figure 2b of the paper cited below shows the geometrical organization of the stacked grana and its description says:

The circular border between the appressed domain and the margin annulus (ring) allows a large contact area between the PSII and the PSI domains and efficient linear electron transport from PSII via plastoquinone (PQ), plastocyanin and cytochrome b6f to PSI at a short distance of ~30–60 nm in local domain.

Further informations can be found in Albertsson, P-Å. (2001) Trends Plant Sci. 6, 349–354.

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