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          are not in direct contact. A small but   to sodium which allows this positive ion to
                                                  follow its concentration and electrical
            distinct separation exists, and the space
  VetBooks.ir  between is the synaptic cleft. Neurotrans­  gradients by flowing to the cell’s interior.
                                                                neurotransmitters
                                                                                   cause
          mitters released by the presynaptic neuron
                                                     Inhibitory
          must diffuse across the synaptic cleft   membrane hyperpolarization by increasing
          to have their effect on postsynaptic cells.   the membrane’s permeability to either
          However, this diffusion occurs almost   potassium (a positive ion that will flow out
          instantaneously because of the very small   of the cell) or chloride (a negative ion that
          size of the cleft (average 20 nm, or about a   will flow into the cell). This makes the cell’s
          millionth of an inch).                  interior more negative, a hyperpolarization
            Neurotransmitters bind to cell mem­   of the cell membrane that therefore reduces
          brane receptors on postsynaptic cells. At   the excitability of the postsynaptic cell.
          synapses between neurons, the binding      The amount of excitatory neurotransmit­
          most commonly changes the postsynaptic   ter released by a single action potential in a
          membrane’s permeability to ions (either   single  presynaptic  neuron is a  constant.
          directly  or indirectly  via second messen­  However, it is also typically insufficient to
          gers), and this in turn produces a change in   depolarize the postsynaptic neuron to the
          the membrane potential of the postsynaptic   threshold voltage at which another action
          neuron. Synapses and the neurotransmit­  potential can be elicited. Therefore, to reach
          ters that depolarize the postsynaptic neuron   threshold voltage in a postsynaptic neuron,
          are  excitatory synapses and  excitatory   the change in voltage produced by multiple
          neurotransmitters, while synapses and   single action potential events must be added
          neurotransmitters that  hyperpolarize the   together, or summated. Summation is classi­
          postsynaptic neuron are inhibitory.     fied as either spatial or temporal (Fig. 11‐6).
            Typically, excitatory neurotransmitters   Spatial summation occurs when simulta­
          cause membrane depolarization by        neous or nearly simultaneous neuro­
          increasing the membrane’s permeability   transmitter release occurs at more than

                            (A)                      (B)




                                         a
                                b



                                                                  d
                                c







                                     Axon                    Axon




          Figure 11-6.  (A) Spatial summation. Each of multiple synapses (a through c) fire simultaneously and
          the additive effects of each synapse brings the postsynaptic neuron to threshold. (B) Temporal summa­
          tion. Repeated, rapid firing of a single synapse (d) allows sufficient depolarization to bring the postsynaptic
          neuron to threshold.
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