Fig. 249The long ascending pathways of the dorsal columns (yellow lines) and spinothalamic tracts (red lines).

Fig. 250The long descending pathway of the pyramidal tract.

Fig. 237The location of the important spinal tracts. (The descending tracts are shown on the left, the ascending tracts on the right.)

Clinical features 1Complete transection of the cord is followed by total loss of sensation in the regions supplied by the cord segments below the level of injury together with flaccid muscle paralysis. As the cord distal to the section recovers from a period of spinal shock, the paralysis becomes spastic, with exaggerated reflexes. Voluntary sphincter control is lost but reflex emptying of bladder and rectum subsequently return, provided that the cord centres situated in the sacral zone of the cord are not destroyed. 2Destruction of the centre of the cord, as occurs in syringomyelia and in some intramedullary tumours, first involves the decussating spinothalamic fibres so that initially there is bilateral loss of pain and temperature sense below the lesion; proprioception and fine touch are preserved till late in the uncrossed posterior columns.

3Hemisection of the cord is followed by the Brown-Squard syndrome; there is paralysis on the affected side below the lesion (pyramidal tract) and also loss of proprioception and fine discrimination (dorsal columns). Pain and temperature senses are lost on the opposite side below the lesion, because the affected spinothalamic tract carries fibres which have decussated below the level of cord hemisection. 4Tabes dorsalis, which is a syphilitic degenerative lesion of the posterior columns and posterior nerve roots, is characterized by loss of proprioception; the patient becomes ataxic, particularly if he closes his eyes, because he has lost his position sense for which he can partially compensate by visual knowledge of his spatial relationship (Rombergs sign). 5Intractable pain can be treated in selected cases by cutting the appropriate posterior nerve roots (posterior rhizotomy) or by division of the spinothalamic tract on the side opposite the pain (cordotomy).

The long ascending and descending pathways The somatic afferent pathways (Fig. 249) 1Proprioceptive and tactile impulses pass uninterruptedly through the posterior root ganglia, through the ipsilateral posterior columns of the spinal cord to the gracile and cuneate nuclei in the lower part of the medulla. In the posterior columns there is a fairly precise organization of the afferent fibres; those from sacral and lumbar segments are situated medially in the tracts while fibres from thoracic and cervical levels are successively added to their lateral aspect. This arrangement according to body segments is maintained in the gracile and cuneate nuclei and in the efferents from these nuclei to the contralateral thalamus. The fibres arising from the gracile and cuneate

nuclei immediately cross over to the opposite side in the sensory decussation of the medulla (Fig. 241) and continue up to the thalamus as a compact contralateral bundlethe medial lemniscus. 2Dorsal root fibres subserving pain and temperature, together with some tactile afferents, end ipsilaterally in the substantia gelatinosa of the posterior horn. They then synapse and cross to the contralateral anterior lateral columns of the cord and are relayed to the contralateral thalamus. The fibre crossing occurs in the anterior white commissure of the spinal cord. In the brainstem these fibres come to lie immediately lateral to the medial lemniscus and are sometimes known as the spinal lemniscus (see Figs 249, 258). They terminate in the thalamus. These somatic afferents are relayed from the thalamus, through the posterior limb of the internal capsule (Fig. 248) to the somatic sensory cortex of the postcentral gyrus. In the internal capsule the fibres are arranged in the sequence face, arm, trunk and leg from before backwards, and this segregation persists in the sensory cortex, where the leg is represented on the dorsal and medial part of the cortex, the trunk and arm in its middle portion and the face most inferiorly. Since the size of the area of cortical representation reflects the density of the peripheral innervation and hence complexity of the function being performed rather than the area of the receptive field, there is a good deal of distortion of the body image in the cortex, the cortical representation of the face and hand being much greater than that of the limbs and trunk.

Clinical features 1Lesions of the sensory pathway most commonly occur in the internal capsule following some form of cerebrovascular accident. If complete, these result in a total hemianaesthesia of the opposite side of the body. In partial lesions the area of sensory loss will be determined by the site of the injury in the internal capsule and, from a knowledge of the sensory (and motor) loss, it is usually possible to determine with some degree of accuracy the site of a lesion in the capsule. 2Since there is modality segregation below the decussation of the medial lemniscus, lesions of the sensory pathways at cord level result in dissociation of sensation, with an area of analgesia contralaterally together with impairment of tactile sensibility ipsilaterally (for further details, see pages 3667). The auditory, visual and olfactory pathways are dealt with later under the appropriate cranial nerves.

The motor pathways (Fig. 250) It is customary to divide the motor pathways of the brain and spinal cord into pyramidal and extrapyramidal systems. Although the latter is an imprecise concept, it provides a useful collective term for the many motor structures not confined to the pyramidal tracts in the medulla.

The pyramidal tract The pyramidal system is the main voluntary motor pathway and derives its name from the fact that projections to the motor neurons in the spinal cord are grouped together in the medullary pyramids. The fibres in this pathway arise from a wide area of the cerebral cortex. About two-thirds

derive from the motor and premotor cortex of the frontal lobes; however, about one-third arises from the primary somatosensory cortex. In both the motor and premotor cortex there is an organization comparable to that seen in the sensory area. Again, the body is inverted so that the leg area is situated in the dorsomedial part of the precentral gyrus encroaching on the medial surface of the hemisphere, supplied by the anterior cerebral artery. The face area is near the lateral sulcus, while the arm area occupies a central position, both supplied by the middle cerebral artery. Again, the body image is greatly distorted; the area representing the hand, lips, eyes and foot are exaggerated out of proportion to the rest of the body and in accordance with the complexity of the tasks they perform. From the cortex, the motor fibres pass through the posterior limb of the internal capsule (Fig. 248) where they are again organized in the sequence of face, arm, leg, anteroposteriorly. From the internal capsule the fibres form a compact bundle which occupies the central third of the cerebral peduncle. Hence they pass through the ventral pons, where they are broken up into a number of small bundles between the cells of the pontine nuclei and the transversely disposed pontocerebellar fibres. Near the lower end of the pons they again collect to form a single bundle which comes to lie on the ventral surface of the medulla and forms the elevation known as the pyramid. As it passes through the brainstem, the pyramidal system gives off, at regular intervals, contributions to the somatic and branchial arch efferent nuclei of the cranial nerves. Most of these corticobulbar fibres cross over in the brainstem, but many of the cranial nerve nuclei are bilaterally innervated. Near the lower end of the medulla the great majority of the pyramidal

tract fibres cross over to the opposite side and come to occupy a central position in the lateral white column of the spinal cord. This is the so-called crossed pyramidal tract shown in Fig. 237. Asmall proportion of the fibres of the medullary pyramid, however, remain uncrossed until they reach the segmental level at which they finally terminate. This is the direct or uncrossed pyramidal tract, which runs downwards close to the anteromedian fissure of the cord, with fibres passing from it at each segment to the opposite side. In view of the frequent involvement of the pyramidal tract in cerebrovascular accidents, its blood supply is listed here in some detail: motor cortex leg area: anterior cerebral artery; face and arm areas: middle cerebral artery; internal capsulebranches of the middle cerebral artery; cerebral peduncleposterior cerebral artery; ponspontine branches of basilar artery; medullaanterior spinal branches of vertebral artery; spinal cordsegmental branches of anterior and posterior spinal arteries. Clinical features 1It is important to remember that, in the motor cortex, movements are represented rather than individual muscles; lesions of this pathway result in paralysis of voluntary movement on the opposite side of the body although the muscles themselves are not paralysed and may cause involuntary movements. This is the essential difference between an upper motor neuron lesion (i.e. a lesion of the central motor pathway) and a lower motor neuron lesion (i.e. a lesion affecting the cranial nerve nuclei, or the

anterior horn cells or their axons). In both types of lesion muscular paralysis results; in the latter, reflex activity is abolished, flaccidity and muscular atrophy follow, whereas, in pyramidal lesions, there is spasticity, increased tendon reflexes and an extensor plantar response. 2Experimental lesions strictly confined to the pyramidal tract are not followed by increased muscular tone in the affected part (spasticity), but clinically this is a feature of upper motor neuron lesions; it is attributable to he central nervous system concomitant involvement of the extrapyramidal system, hence demonstrating the over simplification of the pyramidal and extrapyramidal concept. 3The pyramidal tract is most frequently involved in cerebrovascular accidents where it passes through the internal capsule. Indeed, the artery supplying this area the largest of the perforating branches of the middle cerebral arteryhas been termed the artery of cerebral haemorrhage. 4A list of the more important related signs is given here for involvement of the pyramidal tract at each level. Cortexisolated lesions may occur here, resulting in loss of voluntary movement in, say, only one contralateral limb, but often the sensory cortex is also involved. Aphasia in dominant hemisphere lesions, (usually left), involving Broca and Wernickes areas and the cortex between them, is not uncommon. Internal capsuleusually all parts of the tract are involved, giving a complete contralateral hemiplegia with associated sensory loss. The lesion may extend back to involve the visual radiation, giving a contralateral homonymous field defect (hemianopia).

Cerebral peduncle and midbrainthe fibres from the 3rd nerve are often concomitantly involved so that there are the associated signs of a 3rd nerve palsy. Ponshere the 4th nerve is often involved, alone or together with VII. There may then be a hemiplegia affecting the arm and leg of the opposite side and an abducens and a facial palsy of the lower motor neuron type on the same side as the lesion. Medulla because of the proximity of the pyramids to one another, medullary lesions often affect both sides of the body. Paralysis of the tongue on the side of the lesion is due to involvement of the 12th nerve or its nucleus. The respiratory, vasomotor and swallowing centres may also be affected. Spinal cordthe paralysis following lesions of the spinal cord is ipsilateral and accurately depends on the level at which the pyramidal tract is involved. Lower motor neurone lesion signs can be detected at the level of the spinal trauma (direct injury) and upper motor neurone lesion signs below. The proximity of the pyramidal tracts to the ascending sensory pathways accounts for the concomitant sensory changes which are usually found. The extrapyramidal system The extrapyramidal motor system should, by definition, include all those motor projections which do not pass physically through the medullary pyramids. It was once thought to control movement in parallel with and, to a large extent, independently of the pyramidal motor system and the pyramidal/extrapyramidal division was used clinically to distinguish between two motor syndromes: one characterized by spasticity and paralysis

whereas the other involved involuntary movements, or immobility without paralysis. It is now clear that many extrapyramidal structures, The brain 359 particularly the basal ganglia, actually control movement by altering activity in the premotor cortex and, thus, the pyramidal motor projections. This clearly emphasizes the blur between the two systems. Components of the extrapyramidal system include the red nuclei, vestibular nuclei, superior colliculus and reticular formation in the brain stem, all of which project via discrete pathways to influence spinal cord motor neurons. Cerebellar projections (see page 344) are also included since they influence not only these brainstem motor pathways, but also the motor cortex itself via the dentatothalamic projection. Perhaps the most important structures to retain an extrapyramidal definition are the basal ganglia (see pages 353 and 354). The neostriatum (caudate and putamen) receives widespread cortical afferents, including those from high order sensory association and motor areas, and projects mainly to the globus pallidus. The latter nucleus is the major outflow for the basal ganglia and, via the ventral anterior thalamus, exerts its major influence on premotor and hence the motor cortices. This pattern of connections suggests that the basal ganglia are involved in complex aspects of motor control, including motor planning and the initiation of movement. A variety of motor disorders are associated with basal ganglia pathology and, in some instances, neuroanatomically discrete deficits in specific neurotransmitters. For example, Parkinsons disease involves the degeneration of dopaminergic neurons in the substantia nigra in the midbrain. This pigmented nucleus provides the neostriatum with a dense dopaminergic innervation which may be completely lost in severe cases of Parkinsonism.

Knowledge of this selective chemical neuropathology has resulted in the development of a treatment of the disease which involves the oral administration of the dopamine precursor l-dopa.