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tex to layer V, where these neurons are located, inevitably activating a large number of other cortical neurons The elegant experiments of Asanuma and of Evarts and his colleagues, who stimulated the depths of the cortex with microelectrodes, demonstrated the existence of discrete zones of efferent neurons that control the contraction of individual muscles; moreover, the continued stimulation of a e ttl given efferent zone often facilitated rather than inLi ing le R d x id de b eck hibited the contraction of the antagonists These inM n N s I hum Browl vestigators have also shown that cells in the efferent er al T b ng eye Fi zone receive afferent impulses from the particular nd e d a Fac li Eye muscle to which the efferent neurons project When Lips the effects of many stimulations at various depths Jaw were correlated with the exact sites of each penetraTong Swa ue llow tion, cells that projected to a particular pool of spinal ing motor neurons were found to be arranged in radially ic a st a [M aligned columns about 1 mm in diameter The columnar arrangement of cells in the sensorimotor cortex had been appreciated for many years; the wealth of radial interconnections between the cells in these Lateral Medial columns led Lorente de No to suggest that these Figure 3-4 The representation of body parts in the motor cortex, commonly called the motor vertical chains of cells were the elementary funchomunculus The large area of cortex devoted to motor control of the hand, lips, and face is tional units of the cortex On the sensory side, this evident B in the smaller diagram represents the motor cortex; A is the sensory cortex notion received strong support from Mountcastle s observations that all the neurons in a somatosensory column receive impulses of the same sensory modality, from the separated points From this it may be assumed, as Ash and Georsame part of the body In regard to the functional organization of gopoulus point out, that the premotor cortex includes several anathe columns of motor neurons, it is still not entirely clear whether tomically distinct subregions with different afferent and efferent they contribute to a movement as units or whether individual cells connections In general, it may be said that the motor-premotor within many columns are selectively activated to produce a movecortex is capable of synthesizing agonist actions into an in nite ment The evidence for these disparate views has been summarized variety of nely graded, highly differentiated patterns These are by Henneman and by Asanuma directed by visual (area 7) and tactile (area 5) sensory information The role of cerebrocortical motor neurons in sensory evoked and supported by appropriate postural mechanisms or planned movement has also been elucidated by Evarts and his The supplementary motor area is the most anterior portion of colleagues Using single-cell recording techniques, they showed area 6 on the medial surface of the cerebral hemisphere Stimulation that pyramidal cells re about 60 ms prior to the onset of a moveof this area may induce relatively gross homolateral or contralateral ment in a sequence determined by the required pattern and force movements, bilateral tonic contractions of the limbs, contraversive of the movement But other, more complex properties of the pymovements of the head and eyes with tonic contraction of the conramidal cells were also noted Some of them received a somatotralateral arm, and sometimes inhibition of voluntary motor activity sensory input transcortically from the parietal lobe (areas 3, 1, and and vocal arrest 2), which could be turned on or off or gated according to whether Precisely how the motor cortex controls movements is still a the movement was to be controlled, ie, guided, by sensory input controversial matter The traditional view, based on the interpreMany neurons of the supplementary and premotor parts of the motations of Hughlings Jackson and of Sherrington, is that the motor tor cortical area were activated in a set discharge that preceded cortex is organized not in terms of individual muscles but of move(anticipated) a planned movement Thus, pyramidal (area 4) motor ments, ie, the coordinated contraction of groups of muscles Jackneurons were prepared for the oncoming activation; ie, they were son visualized a widely overlapping representation of muscle triggered by a number of transcortical impulses from the parietal, groups in the cerebral cortex on the basis of his observation that a prefrontal, premotor, and auditory and visual areas of the cortex patient could recover the use of a limb following destruction of the This set signal could occur in the absence of any segmental activity limb area as de ned by cortical stimulation This view was supin the spinal cord and muscles The source of this set signal was ported by Sherrington s observations that stimulation of the cortical found to be mainly in the supplementary motor cortex, which apsurface activated not solitary muscles but a combination of muspears to be under the direct in uence of the readiness stimuli (Bercles, and always in a reciprocal fashion ie, in a manner that eitschaft potential) reaching it from the prefrontal areas (for maintained the expected relationship between agonists and antagplanned movements) and the posterior parietal cortex (for motor onists He also noted the inconstancy of stimulatory effects; the activities initiated by visual, tactile, and auditory perceptions) stimulation of a given cortical point that initiated exion of a part Also, there are bers that reach the motor area from the limbic on one occasion might initiate extension on another system, presumably subserving motivation and attention The These interpretations must be viewed with circumspection, as blood ow to each of the cortical motor zones increases with an must all observations based on the electrical stimulation of the increase in their synaptic activity, and the measurement of blood surface of the cortex It has been shown that to stimulate motor ow has been used to follow these neural events (Roland) cells from the surface, the electric current has to penetrate the corA.

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