Lecture notes: Spinal Cord IV – Sensory Systems & Pathways James Bisley (jbisley@mednet.ucla.edu) Learning objectives: 1. For the 2 main ascending sensory systems, you should know what information they carry, where that information comes from and the pathway it follows to cortex, with a particular emphasis on where information crosses from one side of the CNS to the other. 2. Understand the basic concept underlying the way that pain information can be centrally controlled at the level of the spinal cord. Suggested reading: Purves (4th ed) Chapters 9 & 10. Overview Multiple ascending (sensory) and descending (motor) systems are present within the spinal cord, and each has a distinctive pathway. Knowledge of the pathways is necessary for determining the location of lesions within these systems and for understanding their effects. One general principle is that most motor and sensory pathways cross the midline in some region of the CNS. As a result, one side of the cerebral cortex will be associated with function on the contralateral side of the body. A general rule of thumb is that the sensory pathways cross at the level of the second order neurons’ cell bodies. A second general principle is that there are topographic representations of the body at all levels of the pathways. For instance, in primary somatosensory cortex, information about the lower body is represented medially and information from the head is represented most laterally. Usually the arrangement can be deduced from first principles. Different types of somatosensory information are conveyed by different pathways. Such organization of the somatosensory system has protective value for it helps guard against loss of all sensation following localized damage. In this class, we focus on two distinctive somatosensory systems: the dorsal column-medial lemniscal system and the anterolateral system (frequently identified as the spinothalamic system). The pathways described here and in the lecture represent the main route of information from the periphery to the brain. However, it is important to remember that many of the afferents entering the CNS have collaterals that synapse locally on motor neurons or interneurons. These are often used in reflexes and can also be the start of other ascending pathways, such as in Clarke’s Nucleus. A. Dorsal Column-Medial Lemniscal System This pathway conveys fine touch and position sense from the body to the cerebral cortex. From the body 1. Larger diameter afferents from receptors in the skin, joints and muscles have their cell bodies in the dorsal root ganglia. They enter the dorsal horn through the medial division of the dorsal root and their collaterals proceed to the dorsal columns without a synapse. 2. Afferents from the lower limbs and lower trunk ascend within the medial column (gracile fasciculus or tract) while those from the upper trunk and upper limbs ascend in the lateral column (cuneate fasciculus or tract). 3. The fibers continue in the ipsilateral dorsal column until they reach the lower medulla where they synapse on the second order neurons in the appropriate dorsal column nucleus (gracile nucleus or cuneate nucleus). 4. Axons of these second order neurons curve ventromedially as the internal arcuate fibers and cross the midline of the medulla in the decussation of the medial lemniscus. The fibers are then grouped together in a bundle called the medial lemniscus that is located along the midline of the ventral portion of the medulla. 5. As the medial lemniscus heads rostrally, it rotates so that the topographic representation matches that in the thalamus. 6. The medial lemniscal fibers form synapses on the third order neurons in the ventral posterior lateral (VPL) nucleus of the thalamus. 7. Fibers from the thalamus then project to the primary somatosensory cortex of the parietal lobe (Areas 3a, 3b,1 & 2) via the posterior limb of the internal capsule. Although not passing through the spinal cord, fine touch and position sense from the head and face get processed in a similar way, but in the brain stem. From the face 1. Afferents from receptors in the skin have their cell bodies in the trigeminal ganglion. They then enter the brain stem via the trigeminal nerve. Afferents from receptors in the joints and muscles also enter the brain stem via the trigeminal nerve, but their cell bodies reside within the CNS in the mesencephalic nucleus of the trigeminal complex. 2. Independent of where their cell bodies are located, the afferents all synapse on second order neurons in the sensory or principal nucleus of the trigeminal complex in the pons. 3. Axons from the second order neurons decussate in the pons and join the trigeminothalamic tract, which runs adjacent to the medial lemniscus. 4. Axons in the trigeminothalamic tract form synapses on the third order neurons in the ventral posterior medial (VPM) nucleus of the thalamus. This is medial to the VPL, such that the whole body is represented in the VP complex. 5. Fibers from the thalamus project to the same areas of primary somatosensory cortex. 2 B. Anterolateral System This pathway conveys the sensations of pain, temperature and coarse touch from the periphery to the cortex. From the body 1. Smaller diameter fibers from the skin and deeper structures have their cell bodies in the dorsal root ganglia and enter the spinal cord through the dorsal roots and may ascend or descend 2-3 segments in Lissauer's tract (the dorsolateral tract) before penetrating deeper into the dorsal horn. 2. The fibers then form synapses on the second order neurons in several lamina of the dorsal horn in the spinal cord. 3. The axons of the second order neurons cross the midline of the spinal cord in the anterior white commissure and assemble in the spinothalamic or anterolateral tract on the contralateral side. 4. The fibers of the anterolateral tract remain in lateral locations throughout the brainstem and are eventually located dorsolateral to the medial lemniscus. 5. Upon reaching the thalamus, the axons form synaptic connections with third order neurons in the ventral posterior lateral (VPL) nucleus of the thalamus. Note that in addition, some axons also have synapses in other thalamic nuclei that include the intralaminar and posterior nuclei. 6. Neurons in the ventral posterior lateral nucleus project to the primary somatosensory cortex. Note that on their way through the brainstem, many fibers of the anterolateral system send collaterals into the reticular formation and, by such connections, are believed to influence the level of arousal. Other fibers in this system terminate within the brainstem rather than continuing to the thalamus. Some fibers terminate in the reticular formation and others terminate in the mesencephalon (tectum and periaqueductal gray). Thus, they are referred to as the spinoreticular and spinomesencephalic tracts. As for the dorsal column-medial lemniscal system, there is an equivalent pathway for pain, temperature and coarse touch from the head and face in the brain stem. From the face 1. Afferents have their cell bodies in the trigeminal ganglion and the ganglia associated with the Facial (VII), Glosso-pharyngeal (IX) and Vagus (X) nerves and enter the brain stem via those nerves. 2. Afferents from the trigeminal nerve descend in the spinal trigeminal tract to synapse on second order neurons in the spinal nucleus of the trigeminal complex (primarily the pas caudalis) in the medulla. Afferents from IX and X enter through the medulla and synapse in the same nucleus. 3. Axons from the second order neurons decussate in the medulla and join the anterolateral tract (although one text states that they travel separately from the 3 anterolateral system in the anterior trigeminothalamic tract, we will just refer to it as the anterolateral tract). 4. Fibers in the anterolateral tract containing information from the head and face form synapses on the third order neurons in the ventral posterior medial (VPM) nucleus of the thalamus. Thus, the VP complex contains information from the whole body about all these senses 5. Fibers from the thalamus project to the same areas of primary somatosensory cortex. C. Spinocerebellar Pathways The spinocerebellar tracts will not be traced in detail at this time. However, it is good to appreciate their general organization, i.e. where they are located in the spinal cord and how they reach the cerebellum. The major features of the pathways are outlined below: 1. Both dorsal and ventral spinocerebellar tracts are located in the lateral part of the lateral columns of the spinal cord. 2. These pathways transmit proprioceptive information (non-conscious) from muscle spindles and Golgi tendon organs and some exteroceptive information from cutaneous mechanorecptors to the cerebellum. 3. For both tracts, there are slight differences in the specific pathways for the upper and lower parts of the body. The details of these pathways can be found in the Nolte text. D. Descending pathways for pain modulation While ascending pathways such as the anterolateral system convey pain sensation to higher levels of the nervous system, descending pathways can modulate the sensation of pain. Multisynaptic, descending pathways from the periaqueductal gray in the midbrain to the dorsal horn of the spinal cord (or the spinal nucleus of the trigeminal complex) are capable of decreasing painful sensations. The pathways produce their analgesic effects by inhibiting the nociceptive transmission in the dorsal horn. Local connections from tactile or proprioceptive afferents can also affect transmission of nociceptive information in the dorsal horn. 4