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MUSCLE AND NERVE


noted from it that there is no direct relation between the extent of a cortical area and the mass of muscles which it controls. The mass of muscles in the trunk is greater than in the leg, and in the leg is greater than in the arm, and in the arm is many times greater than in the face and head; yet for the last the cortical area is the most extensive of all, and for the first-named is the least extensive of all.

EB1911 - Volume 19.djvu

Diagram of the Topography of the Main Groups of Foci in the Motor Field
of Chimpanzee.

The motor field of the cortex is, taken altogether, relatively to the size of the lower parts of the brain, larger in the anthropoid than in the inferior monkey brains. But in the anthropoid brain still more increased even than the motor field are the great regions of the cortex outside that field, which yield no definite movements under electric excitation, and are for that reason known as “silent.” The motor field, therefore, though absolutely larger, forms a smaller fraction of the whole cortex of the brain than in the lower forms. The statement that in the anthropoid (orang-outan) brain the groups of foci in the motor fields of the cortex are themselves separated one from another by surrounding inexcitable cortex, has been made and was one of great interest, but has not been confirmed by subsequent observation. That in man the excitable foci of the motor field are islanded inexcitable surface similarly and even more extensively, was a natural inference, but it had its chief basis in the observations on the orang, now known to be erroneous.

In the diagram there is indicated the situation of the cortical centres for movement of the vocal cords. Their situation is at the lower end of the motor field. That they should lie there is interesting, because that place is close to one known in man to be associated with management of the movements concerned in speech. When that area in man is injured, the ability to utter words is impaired. Not that there is paralysis of the muscles of speech, since these muscles can be used perfectly for all acts other than speech. The area in man is known as the motor centre for speech; in most persons it exists only in the left half of the brain and not in the right. In a similar way damage of a certain small portion of the temporal lobe of the brain produces loss of intelligent apprehension of words spoken, although there is no deafness and although words seen are perfectly apprehended. Another region, “the angular region” is similarly related to intelligent apprehension of words seen, though not of words heard.

When this differentiation of cortex, with its highest expression in man, is collated with the development of the cortex as studied in the successive phases' of its growth and ripening in the human infant, a suggestive analogy is obvious. The nervous paths in the brain and cord, as they attain completion, come to be furnished more and more with fibres that are fully myelinate. At the beginning of its history each is unprovided with myelinate nerve fibres. The excitable foci of the cerebral cortex are well myelinated long before the unexcitable are so. The regions of the cortex, whose conduction paths are early completed, may be arranged in groups by their connexions with sense-organs: eye-region, ear-region, skin and somaesthetic region, olfactory and taste region. The areas of intervening cortex, arriving at structural completion later than the above sense-spheres, are called by some association-spheres, to indicate the view that they contain the neural mechanisms of reactions (some have said “ideas”) associated with the sense perceptions elaborated in the several sense-spheres.

The name “motor area” is given to that region of cortex whence, as D. Ferrier’s investigations showed, motor reactions of the facial and limb muscles are regularly and easily evoked. This region is often called the sensori-motor cortex, and the term somaesthetic has Sensori-motor Centres. also been used and seems appropriate. It has been found that disturbance of sensation, as well as disturbance of movement, is often incurred by its injury. Patients in whom, for purposes of diagnosis, it has been electrically excited, describe, as the initial effect of the stimulation, tingling and obscure but locally-limited sensations, referred to the part whose muscles a moment later are thrown into co-ordinate activity. The distinction, therefore, between the movement of the eyeballs, elicited from the occipital (visual) cortex, and that of the hand, elicited from the cortex in the region of the central sulcus (somaesthetic), is not a difference between motor and sensory, for both are sensori-motor in the nature of their reactions; the difference is only a difference between the kind of sense and sense-organ in the two cases, the muscular apparatus in each case being an appanage of the sensual.

That the lower types of vertebrate, such as fish, e.g. carp, possess practically no cortex cerebri, and nevertheless execute “volitional” acts involving high co-ordination and suggesting the possession by them of associative memory, shows that for the existence of these phenomena the cortex cerebri is in them not essential. In the dog it has been proved that after removal from the animal of every vestige of its cortex cerebri, it still executes habitual acts of great motor complexity requiring extraordinarily delicate adjustment of muscular contraction. It can walk, run and feed; such an animal, on wounding its foot, will run on three legs, as will a normal dog under similar mischance. But signs of associative memory are almost, if not entirely, wanting. Throughout three years such a dog failed to learn that the attendant’s lifting it from the cage at a certain hour was the preliminary circumstance of the feeding-hour; yet it did exhibit hunger, and would refuse further food when a sufficiency had been taken. In man, actually gross sensory defects follow even limited lesions of the cortex. Thus the rabbit and the dog are not absolutely blinded by removal of the entire cortex, but in man destruction of the occipital cortex produces total blindness, even to the extent that the pupil of the eye does not respond when light is flashed into the eye.

Examination of the cerebellum by the method of Wallerian degeneration has shown that a large number of spinal and bulbar nerve cells send branches up into it. These seem to end, for the most part, in the grey cortex of the median lobe, some, though not the majority, of them decussating across the median line. The organ seems Cerebellum. also to receive many fibres from the parietal region of the cerebral hemisphere. From the organ there emerge fibres which cross to the opposite red nucleus, and directly or indirectly reach the thalamic region of the crossed hemisphere. The pons or middle, peduncle, which was regarded,