Networking in the endoplasmic reticulum.

The network of the ER (endoplasmic reticulum) is set up by cytoskeletal control of the movement and remodelling of polygonal rings of tubules, bundles of tubules and cisternal regions. We have developed a new image analysis tool, persistency mapping, to understand the framework upon which the plant ER remodels. With this new tool, we have explored the network nodes, called anchor/growth sites, that may anchor the network by attachment to the plasma membrane. We have determined how the polygonal ring structure depends on myosin XI-K for 'opening' and 'closing'. With latrunculin B treatment, we have investigated the involvement of actin in the elongation and persistency of the tubules. We also show how the cytoskeleton is involved in directional diffusion within the membrane. This observation may lead to an answer to the question of what function this network structure serves in the cell. We propose that the ER acts as a trafficking network, delivering lipid, protein, calcium and signalling molecules to different regions of the cell. It does so by directional reduced dimensional diffusion. The ER network of tubules restricts the dimensionality of diffusion to near one-dimensional, whereas the cisternae reduce it to near two-dimensional. The cytoskeleton does not drive the diffusion, but participates by providing directionality to the diffusion.