Corticospinal control during reach, grasp, and precision lift in man

Transcranial magnetic brain stimulation (TMS) was used to assess the influence of the corticospinal system on motor output in seven human subjects during a task in which they had to reach out, grasp, and lift an object. Stimuli, directed at the hand area of the motor cortex, were delivered at eight defined points during the task: during reach, at grip closure, during object manipulation, during the parallel isometric increase in grip and load forces, during the lifting movement, and while the object was held in air. The amplitudes of short-latency EMG responses evoked by TMS in six arm and hand muscles showed a striking modulation across the different phases of the task. This modulation may well reflect phasic changes in corticospinal excitability because: (1) it did not simply reflect phasic changes in muscular activity associated with task performance, (2) it could vary inversely with the amplitude of the background EMG, and (3) it was only obtained with weak TMS intensities, below threshold for evoking responses in hand muscles of the relaxed subject. Our results suggest that the cortical representations of extrinsic hand muscles, which act to orientate the hand and finger tips, were subjected to a strong excitatory drive throughout the reach. This drive was also observed for brachioradialis and anterior deltoid, which contribute to transport of the hand. In contrast, the intrinsic hand muscles appear to receive their strongest cortical input as the digits closed around the object, and just after the subject first touched the object at the onset of manipulation. The isometric parallel increase in load and grip forces necessary to lift the object, which is normally triggered by tactile contact, was delayed by TMS delivered late during the reach. TMS at this time may disrupt processing necessary to control this critical phase of the task.