Philosophical Foundations of Neuroscience. P. M. S. Hacker

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Philosophical Foundations of Neuroscience - P. M. S. Hacker

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the subject destined to dominate his research for over forty years: the mechanism of inhibitory synaptic transmission. After completing his DPhil in 1929, he joined Sherrington’s research group, and developed a technical improvement of the torsion myograph in preparation for a collaboration concerned with research on the flexion reflex and inhibition. These experiments were to see the last flowering of Sherrington’s scientific genius at the age of seventy-five. The work on the ipsilateral spinal flexion reflex introduced Eccles to the technique of stimulating nerves first with just a threshold conditioning volley, then at later intervals with a subsequent test volley in order to tease out the time course of the central excitatory and inhibitory states. This approach, when applied to the mechanism of transmission in the spinal cord, gave a very precise measure of the time course of the central excitatory and inhibitory states, or, as we now know, the excitatory and inhibitory postsynaptic potentials. This was shown by Eccles and his colleagues some twenty years later, when they made the first intracellular recordings of postsynaptic potentials in motoneurones. Subsequent studies of inhibitory synaptic transmission using intracellular electrodes were carried out by Eccles and his colleagues at successively higher levels of the central nervous system. These provided a functional microanatomy of the synaptic connections to be found in the cerebellum, the thalamus and the hippocampus. In this way Eccles completed the research programme described by Sherrington in The Integrative Action of the Nervous System half a century earlier.

      Eccles’s interest in themind–brain problem

      Popper’s influence

      The impact on Eccles of Kornhuber’s research on readiness potential

      Research done by Kornhuber and his colleagues (see §1.6.1) on changes in electrical potential antecedent to a voluntary movement had revealed that the so-called readiness potential began up to 800 milliseconds before the onset of the muscle action potential, and led to a sharper potential, the pre-motion positivity, beginning at 80–90 milliseconds prior to the movement. The patterns of neuronal discharges eventually project to the appropriate pyramidal cells of the motor cortex and synaptically excite them to discharge, so generating the motor potential (a localized negative wave) just preceding the motor pyramidal cell discharge that initiates the movement. The question on which Kornhuber’s research seemed to throw light was: ‘How can willing of a muscular movement set in train neuronal events that lead to the discharge of pyramidal cells of the motor cortex and so to the activation of the neuronal pathways that lead to the muscle contraction?’ (HM 214). It is striking that Eccles took these discoveries to betoken empirical confirmation of mind–brain interaction of a kind (but in a different location) that had been envisaged by Descartes. He argued as follows:

      What is happening in my brain at a time when the willed action is in the process of being carried out? It can be presumed that during the readiness potential there is a developing specificity of the patterned impulse discharges in neurons so that eventually there are activated the pyramidal cells in the correct motor cortical areas for bringing about the required movement. The readiness potential can be regarded as the neuronal counterpart of the voluntary intention. The surprising feature of the readiness potential is its very wide extent and gradual build up. Apparently, at the stage of willing a movement, there is a very wide influence of the self-conscious mind on the pattern of module operation. Eventually this immense neuronal activity is moulded and directed so that it concentrates onto the pyramidal cells in the proper zones of the motor cortex for carrying out the required movement. The duration of the readiness potential indicates that the sequential activity of the large numbers of modules is involved in the long incubation time required for the self-conscious mind to evoke discharges from the motor pyramidal cells … It is a sign that the action of the self-conscious mind on the brain is not of demanding strength. We may regard it as being more tentative and subtle, and as requiring time to build up patterns of activity that may be modified as they develop.(HM 217)

      Cartesian problems recapitulated:

       (1) Interaction

      So, Eccles conceived of what he called ‘the dualist-interactionist hypothesis’ as helping to ‘resolve and redefine the problem of accounting for the long duration of the readiness potential

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