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Chapter 30 – Somatosensory System

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Chapter 30 – Somatosensory System CHAPTER 30 Somatosensory System Jon H. Kaas Vanderbilt University, Nashville, Tennessee, USA OUTLINE Introduction 1075 Ventroposterior Superior Nucleus 1088 Ventroposterior Inferior Nucleus 1088 Receptor Types and Afferent Pathways 1076 The Posterior Group and the Posterior Ventromedial Low-Threshold Mechanoreceptor Afferents Nucleus 1088 from the Hand 1077 Anterior Pulvinar, Medial Pulvinar, and Lateral The SA-I Afferent or the Non-Pacinian Iii Posterior Nucleus 1089 (NP Iii) Channel 1077 The RA-I Afferent or the Non-Pacinian I Anterior Parietal Cortex 1089 (NPI) Channel 1078 Anterior Parietal Cortex in Monkeys 1089 The SA-II Afferent or Non-Pacinian II (NP II) Area 3b 1090 Channel 1078 Area 1 1090 The RA-II Afferent or Pacinian (PC) Channel 1079 Area 2 1091 Cutaneous Receptor Afferents of the Hairy Skin 1079 Area 3a 1091 Deep Receptor Afferents 1080 Subcortical Projections 1091 C-Fiber and A-delta Afferents Mediating Anterior Parietal Cortex in Humans 1091 Temperature, Itch, Pain, and Touch 1080 Architectonic Fields 1091 Afferent Pathways 1081 Maps in 3a, 3b, 1, and 2: Evidence from Scalp Terminations of Peripheral Nerve Afferents in the and Brain Surface Recordings 1092 Spinal Cord and Brainstem 1081 Sensory and Perceptual Impairments Following Ascending Spinal Cord Pathways 1082 Lesions 1094 The Dorsal (Posterior) Column System 1082 Somatosensory Cortex of the Lateral (Sylvian) Second-Order Axons of the Dorsolateral Sulcus Including Insula 1095 Spinal Cord (The Spinomedullothalamic System) 1083 Organization of Cortex of the Lateral Sulcus in The Spinothalamic Pathways 1084 Monkeys 1095 Relay Nuclei of the Medulla and Upper Lateral and Insular Parietal Cortex in Humans 1096 Spinal Cord 1084 Posterior Parietal Cortex 1097 Dorsal ColumneTrigeminal Nuclear Complex 1084 Posterior Parietal Cortex in Monkeys 1097 Lateral Cervical Nucleus, Nuclei X and Z, Area 5a (PE) 1097 External Cuneate Nucleus, and Clarke’s Area 5b (PEc) 1098 Column 1085 Area 7b (PF and PFG) 1098 Lateral Cervical Nucleus 1085 Area 7a (PG) 1098 Nuclei X and Z 1085 Areas of the Intraparietal Sulcus: AIP, LIP, External Cuneate Nucleus 1085 CIP, VIP, MIP, and PIP 1098 Clarke’s Nucleus or Column 1085 Posterior Parietal Cortex Function in Humans 1100 Somatosensory Regions of the Midbrain 1085 Somatosensory Cortex of the Medial Wall: The Somatosensory Thalamus 1086 Supplementary Sensory Area and Cingulate Ventroposterior Nucleus (VP) 1086 Cortex 1101 The Human Nervous System, Third Edition DOI: 10.1016/B978-0-12-374236-0.10030-6 1074 Copyright Ó 2012 Elsevier Inc. All rights reserved. INTRODUCTION 1075 are only briefly mentioned here, as they are reviewed INTRODUCTION elsewhere (e.g., Chapter 32). The basic parts and pathways of the somatosensory This chapter outlines the organization of the somato- system of humans are shown in Figure 30.1. In brief, sensory system of humans. The emphasis is on the peripheral nerve afferents related to receptors in the components of this system that are important in identi- skin, muscles, and joints course centrally past their cell fying objects and features of surfaces by touch. Even bodies in the dorsal root and cranial nerve ganglia to though small shapes can be perceived with information enter the spinal cord and brainstem. These afferents solely from tactile receptors, most discriminations synapse on neurons in the dorsal horn of the spinal involve an active process of tactile exploration and cord, or on equivalent neurons in the brainstem, and multiple contacts on the skin, and an integration of cuta- send a collateral to the dorsal column–trigeminal neous and proprioceptive information as well as efferent nuclear complex. Most of the second-order neurons in control. Thus, this chapter concentrates on the pathways this complex have axons that cross to the contralateral and neural centers for processing information from the lower brainstem to form the medial lemniscus column low-threshold mechanoreceptors of the skin that of axons that ascend to the somatosensory thalamus. provide information about touch, and the deeper recep- Second-order neurons representing the teeth and tongue tors in joints and muscles that provide information project both ipsilaterally and contralaterally to the thal- about position. Conclusions are based on both studies amus (see Chapter 32). Other second-order neurons in humans and studies in other primates, especially in the dorsal horn of the spinal cord and in the brainstem the frequently studied macaque monkeys. At least the send axons to the contralateral side to form the early stages of processing are likely to be similar in anterolateral spinothalamic ascending pathway (see humans and monkeys, but humans appear to have Figure 30.4). The low-threshold mechanoreceptor infor- a more expanded cortical network for processing mation from the skin and the information from muscle somatosensory information. The important subsystems spindle receptors and joints course in the medial dealing with afferents coding for pain and temperature lemniscus to terminate in nuclei of the ventroposterior FIGURE 30. 1 The basic components of the somatosensory system shown on a posterolateral view of a human brain. Receptors in the skin project to the dorsal column nuclei or the trigeminal nuclei to form the dorsal column trigeminal complex. The trigeminal complex in the brainstem provide second-order afferents, representing the face, that join those from the dorsal column nuclei, representing the body, to terminate in the contralateral ventroposterior complex of the thalamus. Third-order neurons in the thalamus project to anterior parietal cortex, where information is distributed to posterior and lateral parietal cortex. VI. SYSTEMS 1076 30. SOMATOSENSORY SYSTEM thalamic complex. The spinothalamic pathway carries VPI and Pa project widely to areas of somatosensory information about pain, temperature, and touch to infe- cortex, and LP projects to posterior parietal cortex. Areas rior and caudal parts of the ventroposterior complex. 3b, l, and 2 form a hierarchical sequence of processing, The somatosensory thalamus has been subdivided in while area 3a relates especially to motor cortex. All various ways (e.g., Chapter 19; Hirai and Jones, 1989; four areas project to somatosensory cortex of the lateral Mai et al., 1997; Morel et al., 1997; Jones, 2007). Because sulcus (lateral parietal cortex), especially the second some of the earlier parcellations of thalamic nuclei were somatosensory area, S2, and a more recently discovered based solely in cytoarchitectonic and myeloarchitectonic parietal ventral somatosensory area, PV (Krubitzer et al., criteria (e.g., Ha¨ssler, 1959), and because they do not 1995). These and other areas of the lateral sulcus relate to reflect the many recent advances in understanding motor areas of cortex to guide action, and to perirhinal based on microelectrode recordings, studies of connec- cortex to engage the hippocampus in object recognition tions, and chemoarchitecture, we use more recently and memory (Mishkin, 1979). Other connections of area proposed subdivisions stemming from studies largely 2 and subdivisions of lateral parietal cortex are with in monkeys (see Kaas, 2008 for review). Thus, the relay subdivisions of posterior parietal cortex that are of afferents from the skin via the medial lemniscus is involved with the early stages of forming movement to the ventroposterior nucleus (VP), which has been intentions and sensory guidance for motor actions (Des- traditionally subdivided into a ventroposterior medial murget et al., 2009). Subdivisions of the posterior pari- subnucleus (VPM) representing the face and a ventro- etal cortex project to premotor and motor areas of the posterior lateral subnucleus (VPL) representing the frontal lobe (e.g., Tanne-Gariepy et al., 2002; Stepniew- body. The muscle spindle receptor inputs are relayed ska et al., 2009b). Projections from the posterior parietal to a dorsal or superior part of the ventroposterior cortex also involve the medial limbic cortex of the pari- complex that is often included in VPM and VPL. We etal lobe in producing motivational and possibly distinguish this representation of deep body receptors emotional states. as the ventroposterior superior nucleus (VPS). Many of the spinothalamic afferents terminate in a ventroposte- rior inferior nucleus (VPI). Other spinothalamic affer- RECEPTOR TYPES AND AFFERENT ents end in a caudal part of the VP complex that is PATHWAYS often included in VPI or end in the posterior group of nuclei, part of which has been distinguished as a distinct In humans, the hand is the most important tactile relay nucleus for pain and temperature, the ventrome- organ for object identification (Darian-Smith, 1984). dial posterior nucleus (VMpo) (Dostrovsky and Craig, Receptors in the hand must convey information about 2008). The somatosensory thalamus also includes nuclei texture and shape. This is done primarily from the finger without ascending somatosensory inputs. The anterior pads during active exploration; consequently, finger pulvinar (Pa) and the lateral posterior complex (LP) position and temporal sequence are important. The receive inputs from somatosensory cortex and project glabrous (hairless) skin of the hand has the highest back to somatosensory cortex. innervation density and tactile acuity of any body The somatosensory cortex has also been variously surface (Darian-Smith, 1984). Hairy skin is less impor- subdivided into proposed subdivisions of functional tant in object identification, and hairy skin
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