Membrane Electrochemistry and Bioelectrochemistry in .NET

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6 Membrane Electrochemistry and Bioelectrochemistry
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Not only electric currents in muscles and nerves but particularly even the mysterious effects occurring with electric fish can be explained by means of the properties of semipermeable membranes. W. Ostwald, 1891 6.1 Basic Concepts and Definitions
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The discovery of galvanic electricity (i.e. electrical phenomena connected with the passage of electric current) by L. Galvani in 1786 occurred simultaneously with his study of a bioelectrochemical phenomenon which was the response of excitable tissue to an electric impulse. E. du Bois-Reymond found in 1849 that such electrical phenomena occur at the surface of the tissue, but it was not until almost half a century later that W. Ostwald demonstrated that the site of these processes are electrochemical semipermeable membranes. In the next decade, research on semipermeable membranes progressed in two directions in the search for models of biological membranes and in the study of actual biological membranes. The search for models of biological membranes led to the formation of a separate branch of electrochemistry, i.e. membrane electrochemistry. The most important results obtained in this field include the theory and application of ion-exchanger membranes and the discovery of ion-selective electrodes (including glass electrodes) and bilayer lipid membranes. The study of biological membranes led to the conclusion that the great majority of the processes in biological systems occur at cell and organelle membranes. The electrochemical aspects of this subject form the basis of bioelectrochemistry, dealing with the processes of charge separation and transport in biological membranes and their models, including electron and proton transfer in cell respiration and photosynthesis as well as ion transport in the channels of excitable cells. The electrokinetic phenomena (electrical double layer, interfacial tension of cells and organelles, cell membrane extension and contraction, etc.) also belong to this field. Bioelectrochem410
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411 istry includes the more classical subjects of the thermodynamics and kinetics of redox processes of components of biological systems at electrodes. This chapter will deal with the basic properties of electrochemical membranes in general and the membrane aspects of bioelectrochemistry in particular. A number of bioelectrochemical topics was discussed in Sections 1.5.3 and 3.2.5. 6.1.1 Classification of membranes In contrast to mechanics, where the term membrane (Lat. membrana = parchment) designates an elastic, two-dimensional plate, this term is used in chemistry, biophysics and biology to designate a solid or liquid phase (usually, but not always, with a thickness substantially smaller than its other dimensions) separating two, usually liquid, phases. The transport (permeation) of the various components of both phases through the membrane occurs at different rates relative to those in the homogeneous phases with which the membrane is in contact. The membrane is consequently called semipermeable. The thickness of the membrane phase can be either macroscopic ('thick') membranes with a thickness greater than micrometres or microscopic ('thin'), i.e. with thicknesses comparable to molecular dimensions (biological membranes and their models, bilayer lipid films). Thick membranes are crystalline, glassy or liquid, while thin membranes possess the properties of liquid crystals (fluid) or gels (crystalline). Depending on their structure, membranes can be separated into porous, where matter is transported through pores in the membrane, and compact, where the substance is transported either through the entire homogeneous membrane phase or its homogeneous parts. Membranes can be homogeneous, where the whole membrane participates in the permeation of a substance, or heterogeneous, where the active component is anchored in a suitable support (for solid membranes) or absorbed in a suitable diaphragm or acts as a plasticizer in a polymeric film. Both of the latter cases are connected with liquid membranes. Biological membranes show heterogeneity at a molecular level. Membranes exhibiting selectivity for ion permeation are termed electrochemical membranes. These membranes must be distinguished from simple liquid junctions that are often formed in porous diaphragms (see Section 2.5.3) where they only prevent mixing of the two solutions by convection and have no effect on the mobility of the transported ions. It will be seen in Sections 6.2 and 6.3 that the interior of some thick membranes has properties analogous to those of liquid junctions, but that the mobilities of the transported ions are changed. 6.1.2 Membrane potentials A characteristic property of electrochemical membranes is the formation of an electric potential difference between the two sides of the membrane,
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412 termed the membrane potential A$ M : d <H2) solution 1 | membrane | solution 2 (6.1.1) x=p x=q Similar to galvanic cells, the membrane potential is determined by subtracting the electric potential of the phase on the left from that of the phase on the right, i.e.
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A0 M -A 0 = <K2)-<H1)
(6.1.2)
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For cell membranes, the intracellular liquid is usually denoted as solution 2, while solution 1 is the extracellular liquid. The formation of a membrane potential is connected with the presence of an electrical double layer at the surface of the membrane. For a thick, compact membrane, an electrical double layer is formed at both interfaces. The electrical double layer at a porous membrane is formed primarily in the membrane pores (see Section 6.2). The electrical double layer at thin membranes is formed on both membrane surfaces. It is formed by fixed ions on the surface of the membrane and the diffuse layer in the electrolyte. Consider the simple case where both sides of the membrane are in contact with a solution of symmetrical electrolyte BA in a single solvent and the membrane is permeable for only one ionic species. In equilibrium its electrochemical potential (Eq. (3.1.5)) in both solutions adjacent to the membrane has the same value. Thus,
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