Physiological studies of electric and magnetic stimulation of the human brain.

It is suggested that transcranial electric stimulation can activate pyramidal tract projections both directly and indirectly in a manner similar to that described after direct stimulation of the exposed cortex in the monkey. This produces both D- and I-waves in the pyramidal tract. At high intensities of stimulation, the stimulus can spread into the brain and activate pyramidal tract axons several centimeters below the cortical surface. Magnetic stimulation at moderate intensities produces electromyographic (EMG) responses with latencies 1-2 msec longer than those after electric stimulation. Two possible explanations have been put forward to account for this effect: (1) because of the difference in the direction of electric current flow induced in the brain by the 2 forms of stimulation, magnetic stimulation preferentially excites pyramidal tract cells indirectly, and hence evokes only I-waves in the pyramidal tract. (2) Electric stimulation (even at threshold) activates pyramidal axons deep in the white matter, whereas magnetic stimulation activates the pyramidal cells in the gray matter, probably at their initial segment. There is one interesting consequence common to both explanations. Whether magnetic stimulation activates the pyramidal neurons transsynaptically or at their initial segment, the size of the descending volley evoked will depend on the level of excitability of the motor cortex. In contrast, the response to electric stimulation will be less affected, since a proportion of the descending volley is initiated directly at the axon of the pyramidal cell. This differential effect of cortical excitability on the responses to electrical and magnetic stimulation can be useful in describing excitatory or inhibitory influences on motor cortex from other structures.