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17 the nerve cell is linked to other excitable, both nerve and muscle, cells by structures called, in the case of other nerve cells, as partners, synapses, and in the case of striated muscle cells, motor endplates (neuromuscular junctions) The impulse, which is originally electric, is transformed into a chemical stimulus and again into an electrical impulse The opening and closing of ion-selective channels present in these junctions depend on either electric or chemical actions The substances that are active in the latter case are called neurotransmitters A very important member of this family is acetylcholine which is transferred to the cell that receives the signal across the postsynaptic membrane or motor endplate through a.

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Sgn()

Sgn( number)

Fig. 6.24 A hypothetic scheme of the time behaviour of the spike linked to the opening and closing of sodium and potassium channels. After longer time intervals a temporary hyperpolarization of the membrane is induced by reversed transport of potassium ions inside the nerve cell. Nernst potentials for Na+ and K+ are also indicated in the figure. (According to A. L. Hodgkin and A. F. Huxley)

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specific channel, the nicotinic acetylcholine receptor. This channel has been investigated in detail, because, among other reasons, it can be isolated in considerable quantities as the membranes of the cells forming the electric organ of electric fish are filled with this species. Acetylcholine, which is set free from vesicles present in the neighbourhood of the presynaptic membrane, is transferred into the recipient cell through this channel (Fig. 6.25). Once transferred it stimulates generation of a spike at the membrane of the recipient cell. The action of acetylcholine is inhibited by the enzyme, acetylcholinesterase, which splits acetylcholine to choline and acetic acid. Locomotion in higher organisms and other mechanical actions are made possible by the striated skeletal muscles. The basic structural unit of muscles is the muscle cell muscle fibre which is enclosed by a sarcoplasmatic membrane. This membrane invaginates into the interior of the fibre through transversal tubules which are filled with the intercellular liquid. The inside of the fibre consists of the actual sarcoplasm with inserted mitochondria, sarcoplasmatic reticulum and minute fibres called myofibrils, which are the organs of muscle contraction and relaxation. The membrane of the sarcoplasmatic reticulum contains Ca2+-ATPase, maintaining a concentration of calcium ions in the sarcoplasm of the relaxed muscle below 10~7mol dm~3. Under these conditions, the proteins, actin and myosin, forming the myofibrils lie in relative positions such that the muscle is relaxed. When a spike is transferred from the nerve fibre to the sarcoplasmatic membrane, another spike is also formed there which continues

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BitAnd()

BitAnd( number1, number2)

BitMaskClear()

Fig. 6.25 The nicotinic acetylcholine receptor in a membrane. The deciphering of the structure is based on X-ray diffraction and electron microscopy. (According to Kistler and coworkers) through the transverse tubules to the membrane of the sarcoplasmatic reticulum, increasing the permeability of the membrane for calcium ions by five orders within 1 millisecond, so that the concentration of Ca2+ ions in the sarcoplasm increases above 10~3 mol dm~3. This produces a relative shift of actin and myosin molecules and contraction of the muscle fibre. After disappearance of the spike, Ca2+-ATPase renews the original situation and the muscle is relaxed. 6.5.2 Membrane principles of bioenergetics The basic process by which living organisms obtain energy is photosynthesis in green plants, yielding carbonaceous substances (fuels) and oxidative phosphorylation in all organisms, in which the Gibbs energy obtained by oxidation of these carbonaceous substances with oxygen is converted into the energy of the polyphosphate bond in adenosinetriphosphate (ATP) through the reaction of adenosine diphosphate with dihydrogen phosphate ions (PinOrg)This process occurs in the mitochondria, organelles present in the cells of all multicellular organisms (see Figs 6.8 and 6.26). Mitochondria have two membranes. The invaginations of the internal membrane into the inner space of the organelle (matrix space) are termed crests (from the Latin, cristae). The oxidation of the carbonaceous substrate with oxygen cannot occur directly as the released energy would be converted into heat, but must proceed almost reversibly, through a number of poorly water-soluble redox

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