Genes, Morphogenesis, Evolution: Life and ALife Aspects

Excitable Cells
ELECTRICAL ACTIVITY OF BIOMEMBRANES AND VECTORIZATION OF INTRACELLULAR PROCESSES

V.A.Kaimanovich (1), E.M.Krupitski (2) and A.V.Spirov (3)
1) Department of Mathematics and Statistics, University of Edinburgh, Edinburgh EH9 3JZ, Great Britain
2) Research Laboratory of the Regional Narcological Center, St.Petersburg, Russia
3) I.M. Sechenov Institute of Evolutionary Physiology & Biochemistry St.Petersburg, Russia

DISCUSSION

Directionalized transport of macromolecules (including mRNAs) in subcortical layers of cytoplasm and into plasmalemma of animal and vegetal poles can be carried out both due to intracellular electrophoresis and due to transport along MT bundles oriented from the nuclear envelope to wards the inner surface of plasmalemma (5). Processes of this kind can provide a basis of MT-dependent molecular differentiation of the zygote cortex observed in the developmental biology in various situations.

According to Ysraeli with collaborators (6), the simplest model to explain these results is the two-step process with MT involved in the translocation process and microflaments involved in anchoring the message at the cortex. The apparent involvement of MT in the translocation of the message may correspond to active movement of the message. A number of different types of cells make use of MT-mediated motors to actively direct the transport of macromolecules and organelles. Particularly , MT mediate the localization of bicoid RNA during the Drosophila oogenesis (8).

MT arrays begin to form early in oogenesis; tubulin staining appears frst perinuclearly and then in radial arrays extending completely around the oocyte from the germinal vesicle to the cortex. As oogenesis proceeds and the germinal vesicle migrates to the animal hemisphere, the MT arrays in the vegetal hemisphere remain radially aligned, emanating from the germinal vesicle to the animal cortex (31).

Thus, a hypothetical picture of the intermediate filament net work in cells: vimentin- or desmin-containing filaments associate with the nuclear lamina through the nuclear pores. At the cell periphery the same intermediate filaments apparently associate with the membrane skeleton, thus forming an elaborate system of nucleolemmal/ plasmalemmal interactions. These mechanical interconnections between the nucleus and the membrane can play a general role in the spatial organization of the cytoplasmic matrix or in the active or passive transport of macromolecules between nucleoplasmic and cytoplasmic compartments (31).

A significant role should belong to dynamical regimes of electrically active reaction-diffusion membrane systems. This is especially important for oocytes and zygotes. The plasmalemma of these cells is considered to be excitable (32). There are facts confirming complicated electrical activity of oocytes and other cells (33).

This leads to a conclusion that one of the sources of the information which orders vector processes in the cytoplasm is provided by electrically active dissipative structures in membranes interacting with the cortical part of cytoskeleton. Signals propagating both from the inside and the outside of the nucleus cause transformations of the pattern of electrical fields and ionic currents in the cell and, as a consequence, changes of the cytoskeleton which bring about changes of directed movements of many cytoplasmic components. This mechanism can serve as a foundation of morphogenetic processes in embryogenesis of animals and plants, because these processes are based on directionalized mitosis, directionalized movements, directionalized growth and polarization of cells, vector character of which is intimately related to MT.

In neurons MT are involved not only in transport processes. More and more data about the role of MT in the processes of nervous impulse transmitting is being collected (34, 35). MT take part in processes of synapses modification (36) which are related to the mechanisms of long-term memory . These processes can also be based on the directionalization of MT assemblage by intracellular electric fields. These suggestions seem to be especially topical in connection with a well known idea that local changes in the neuronal membrane play a fundamental role in the mechanisms of memory (37, 38). It is shown, in particular, that these changes originate at a neuron's receiving site when an input there from a conditioned stimulus is temporally associated with the input from an unconditioned stimulus at adjacent receiving site. The interaction of both input sites is communicated to the cell nucleus, generating factors which when directionally transported return to the site of the interaction and "hard-wire" it (38, 39).

Thus, the suggested possible mechanism of MT polymerization by intracellular electric fields and organization of cytoskeleton by electrically active membrane dissipative structures can be also essential for such phenomena as morphogenesis and functioning of the nerve system.

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