The structural organization of layer IV in the somatosensory region (SI) of mouse cerebral cortex. The description of a cortical field composed of discrete cytoarchitectonic units.

The involvement of the basal ganglia in motor functions has been well studied. Recent neurophysiological, clinical and behavioral experiments indicate that the basal ganglia also process non-noxious and noxious somatosensory information. However, the functional significance of somatosensory information processing within the basal ganglia is not well understood. This review explores the role of the striatum, globus pallidus and substantia nigra in nociceptive sensorimotor integration and suggests several roles of these basal ganglia structures in nociception and pain. Electrophysiological experiments have detailed the non-nociceptive and nociceptive response properties of basal ganglia neurons. Most studies agree that some neurons within the basal ganglia encode stimulus intensity. However, these neurons do not appear to encode stimulus location since the receptive fields of these cells are large. Many basal ganglia neurons responsive to somatosensory stimulation are activated exclusively or differentially by noxious stimulation. Indirect techniques used to measure neuronal activity (i.e., positron emission tomography and 2-deoxyglucose methods) also indicate that the basal ganglia are activated differentially by noxious stimulation. Neuroanatomical experiments suggest several pathways by which nociceptive information may reach the basal ganglia. Neuroanatomical studies have also indicated that the basal ganglia are rich in many different neuroactive chemicals that may be involved in the modulation of nociceptive information. Microinjection of opiates, dopamine and gamma-aminobutyric acid (GABA) into the basal ganglia have varied effects on pain behavior. Administration of these neurochemicals into the basal ganglia affects supraspinal pain behaviors more consistently than spinal reflexive behaviors. The reduction of pain behavior following electrical stimulation of the substantia nigra and caudate nucleus provides additional evidence for a role of the basal ganglia in pain modulation. Some patients with basal ganglia disease (e.g., Parkinson's disease, Huntington's disease) have alterations in pain sensation in addition to motor abnormalities. Frequently, these patients have intermittent pain that is difficult to localize. Collectively, these data suggest that the basal ganglia may be involved in the (1) sensory-discriminative dimension of pain, (2) affective dimension of pain, (3) cognitive dimension of pain, (4) modulation of nociceptive information and (5) sensory gating of nociceptive information to higher motor areas. Further experiments that correlate neuronal discharge activity with stimulus intensity and escape behavior in operantly conditioned animals are necessary to fully understand how the basal ganglia are involved in nociceptive sensorimotor integration.

The role of the basal ganglia in nociception and pain

The posterior nuclear complex of the thalamus in rhesus, pigtailed and squirrel monkeys consists of the combined suprageniculate-limitans nucleus and an ill defined region of heterogeneous cell types extending anteriorly from the dorsal lobe of the medial geniculate body towards the posterior pole of the ventral nuclear complex. This region is referred to as the posterior nucleus. It is directly continuous with the ventroposteroinferior nucleus. The cortical projections of each of these nuclei, together with those of the adjacent ventral, pulvinar and medial geniculate complexes, have been studied by means of the autoradiographic tracing technique. The suprageniculate-limitans nucleus, the main input to which is the superior colliculus, projects upon the granular insular area of the cortex. The medial portion of the posterior nucleus projects to the retroinsular field lying posterior to the second somatic sensory area. There is clinical and electrophysiological evidence to suggest that the retroinsular area may form part of a central pain pathway. The lateral portion of the posterior nucleus which is closely related to certain elements of the medial geniculate complex, projects to the postauditory cortical field. The ventroposterioinferior nucleus, which may be involved in vestibular function, projects to the dysgranular insular field. The principal medial geniculate nucleus can be subdivided into a ventral division that projects to field AI of the auditory cortex and a dorsal division that merges with the posterior nucleus; it is further subdivided into an anterodorsal component that projects to two fields on the superior temporal gyrus, together with a posterodorsal component in which separate cell populations project to areas lying anterior and medial to AI. The magnocellular medial geniculate nucleus, sometimes considered a part of the posterior complex, appears to project diffusely to layer I of all the auditory fields. The auditory fields are bounded on three sides by the projection field of the medial nucleus of the pulvinar which also extends into the upper end of the lateral sulcus to bound the fields receiving fibers from the posterior nucleus. The topography of the areas receiving fibers from the posterior, medial geniculate and pulvinar complexes, taken in conjunction with the rotation of the primate temporal lobe, permits all of these fields to be compared with similar, better known areas in the cat brain.

The posterior thalamic region and its cortical projection in new world and old world monkeys

The cytoarchitecture of layer IV of SmI neocortex of the rat has been studied in sections parallel and perpendicular to layer IV. The neurons of layer IV are arranged into discrete multicellular units, called barrels, which we have previously described in both mouse and rat. Using the barrels as markers, it is possible to outline precisely a cytoarchitectonic field—the barrel field—which (1) is confined to SmI. (2) has a consistent appearance from hemisphere to hemisphere, and (3) contains at least 220 barrels. A special region of the barrel field has been identified—the posteromedial barrel subfield (PMBSF)—in which the barrels are arranged in five rows and are larger than elsewhere. However, in contrast to other barrels in the rat barrel field and to the barrels that make up the PMBSF in the mouse, the PMBSF barrels in the rat are filled with small neurons nearly throughout the thickness of layer IV.



From experimental evidence, it has been possible for the first time to establish consistent homologies between small groups of neocortical neurons in layer IV. The most surprising finding is that homologous barrels in the PMBSF of two closely related species, the rat and the mouse, have such a different cytoarchitectonic appearance. On the basis of this observation, we suggest that a careful study of these PMBSF barrels in the mouse, the rat and other species may provide a clue to understanding how neocortical neuronal circuits of varying complexity are assembled.

S. A. Andersson

Papers

Functional properties of neurons of the anterior ectosylvian gyrus of the cat.