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The Electroencephalogram (EEG)
       The neurons of the cerebral cortex, when their  fibers are more likely to lead to positive poten-
       membrane potential is changed, generate  tials at the surface (→ A2) because they act
       varying electrical fields on the surface of the  near the cell body, i.e., deep in the cerebral cor-
       skull that can be recorded with suitable leads.  tex. Inhibition in the area of the cell body the-
       The EEG can provide valuable clues to neuro-  oretically results in a negative potential at the
    Systems  nal functions and as a result has gained great  surface, but it is not strong enough to be regis-
       clinical importance. Like the electrocardio-
                                       tered at the surface of the scalp (→ A3).
                                        The neurons in the thalamus that excite the
       gram ([ECG] → p.184), the EEG registers the
                                       cortical pyramidal cells undergo a rhythmical
       summated activity of the cells that, projected
    Neuromuscular and Sensory  similarly directed dipoles.  rhythm is transmitted by the thalamocortical
                                       activity due to negative feedback (→ A4). This
       onto the area of the recording lead, generates
                                       tracts to the pyramidal cells, with one thalamic
         The potential changes on the cortical sur-
                                       neuron simultaneously exciting several py-
       face largely depend on the postsynaptic poten-
       tials at dendrites of the pyramidal cells (→ A).
                                       ramidal cells. Because of this, subcortical le-
       Although the postsynaptic potentials have a
                                       sions are better registered in the EEG than
                                       small cortical ones.
       lower amplitude than the action potentials,
       they last significantly longer. Because the py-
                                        The frequency of the recorded waves (de-
       the cortical surface, their local activity gener-
                                       on when analysing the EEG (→ B1). In adults
                                       who are awake with their eyes open it is pre-
       ates dipoles in the direction of the surface
    10  ramidal cells are positioned at right angles to  flections) is a diagnostically significant criteri-
                                       dominantly β-waves (14–30 Hz) that are reg-
       much more easily than other cells in the cor-
       tex. They thus have a much greater impact on  istered. With their eyes closed the somewhat
       the surface potential than other neurons. Fur-  slower α-waves (8–3 Hz) dominate. Yet slower
       thermore, they are all orientated in parallel to  waves such as the ϑ-waves (4–7 Hz) and the δ-
       one another, so that equidirectional potential  waves (0.5–3 Hz) are not normally recorded in
       changes of neighboring pyramidal cells are  waking adults but only in children and adoles-
       summated. EEG deflections are to be expected  cents. However, in adults the latter slow waves
       only if (around the lead electrode) several py-  are recorded during the phases of deep sleep
       ramidal cells are simultaneously depolarized,  (→ p. 340). Some diseases of the brain can re-
       i.e., there is a synchronized event.  sult in slowing (sleeping-drug overdose, de-
         During an excitatory postsynaptic potential,  mentia, schizophrenia) or acceleration (alco-
         +
       Na enters the cell and thus leaves behind a lo-  holism, manic-depressive illness) of the re-
       cal negative extracellular potential (→ A1).  corded frequency.
       The depolarization promotes an efflux of K +  The EEG is of particular importance when
       ions along the remaining cell membrane, this  diagnosing epilepsy, which is characterized by
       efflux in turn generating a local positive extra-  massive synchronized excitation of cortical
       cellular potential. If an excitatory synapse at  neurons (→ p. 338). It causes “spike” activity
       the apical end of a dendrite is activated, the ex-  (“seizure spikes”; → B2) or “spike and wave”
       tracellular space in the area is relatively nega-  complexes (→ B3).
       tive, but relatively positive at the base of the  In destruction of the cerebral cortex (brain
                                 +
       dendrite (→ A1; to simplify matters the K ef-  death) all electrical activity will have ceased
       flux has been entered at only one site). As a re-  and the EEG tracing will therefore be isoelec-
       sult a dipole is generated that creates a nega-  tric (“flat”), i.e., there will be no deflections.
       tive potential at the surface. Commissural fi-
       bers from the other cortical hemisphere and
       nonspecific parts of the thalamus form excit-
       atory synapses mainly at the surface; excit-
       ation via these fibers thus leads to a negative
  336  potential at the surface electrode (→ A1). Con-
       versely, activation of specific thalamocortical
       Silbernagl/Lang, Color Atlas of Pathophysiology © 2000 Thieme
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