Pain. Part 2a: Trigeminal Anatomy Related to Pain.

In order to understand the underlying principles of orofacial pain it is important to understand the corresponding anatomy and mechanisms. Paper 1 of this series explains the central nervous and peripheral nervous systems relating to pain. The trigeminal nerve is the 'great protector' of the most important region of our body. It is the largest sensory nerve of the body and over half of the sensory cortex is responsive to any stimulation within this system. This nerve is the main sensory system of the branchial arches and underpins the protection of the brain, sight, smell, airway, hearing and taste, underpinning our very existence. The brain reaction to pain within the trigeminal system has a significant and larger reaction to the threat of, and actual, pain compared with other sensory nerves. We are physiologically wired to run when threatened with pain in the trigeminal region and it is a 'miracle' that patients volunteer to sit in a dental chair and undergo dental treatment. Clinical Relevance: This paper aims to provide the dental and medical teams with a review of the trigeminal anatomy of pain and the principles of pain assessment.

Bilateral descending hypothalamic projections to the spinal trigeminal nucleus caudalis in rats.

Several lines of evidence suggest that the hypothalamus is involved in trigeminal pain processing. However, the organization of descending hypothalamic projections to the spinal trigeminal nucleus caudalis (Sp5C) remains poorly understood. Microinjections of the retrograde tracer, fluorogold (FG), into the Sp5C, in rats, reveal that five hypothalamic nuclei project to the Sp5C: the paraventricular nucleus, the lateral hypothalamic area, the perifornical hypothalamic area, the A11 nucleus and the retrochiasmatic area. Descending hypothalamic projections to the Sp5C are bilateral, except those from the paraventricular nucleus which exhibit a clear ipsilateral predominance. Moreover, the density of retrogradely FG-labeled neurons in the hypothalamus varies according to the dorso-ventral localization of the Sp5C injection site. There are much more labeled neurons after injections into the ventrolateral part of the Sp5C (where ophthalmic afferents project) than after injections into its dorsomedial or intermediate parts (where mandibular and maxillary afferents, respectively, project). These results demonstrate that the organization of descending hypothalamic projections to the spinal dorsal horn and Sp5C are different. Whereas the former are ipsilateral, the latter are bilateral. Moreover, hypothalamic projections to the Sp5C display somatotopy, suggesting that these projections are preferentially involved in the processing of meningeal and cutaneous inputs from the ophthalmic branch of the trigeminal nerve in rats. Therefore, our results suggest that the control of trigeminal and spinal dorsal horn processing of nociceptive information by hypothalamic neurons is different and raise the question of the role of bilateral, rather than unilateral, hypothalamic control.

Corneal afferents differentially target thalamic- and parabrachial-projecting neurons in spinal trigeminal nucleus caudalis.

Dorsal horn neurons send ascending projections to both thalamic nuclei and parabrachial nuclei; these pathways are thought to be critical pathways for central processing of nociceptive information. Afferents from the corneal surface of the eye mediate nociception from this tissue which is susceptible to clinically important pain syndromes. This study examined corneal afferents to the trigeminal dorsal horn and compared inputs to thalamic- and parabrachial-projecting neurons. We used anterograde tracing with cholera toxin B subunit to identify corneal afferent projections to trigeminal dorsal horn, and the retrograde tracer FluoroGold to identify projection neurons. Studies were conducted in adult male Sprague-Dawley rats. Our analysis was conducted at two distinct levels of the trigeminal nucleus caudalis (Vc) which receive corneal afferent projections. We found that corneal afferents project more densely to the rostral pole of Vc than the caudal pole. We also quantified the number of thalamic- and parabrachial-projecting neurons in the regions of Vc that receive corneal afferents. Corneal afferent inputs to both groups of projection neurons were also more abundant in the rostral pole of Vc. Finally, by comparing the frequency of corneal afferent appositions to thalamic- versus parabrachial-projecting neurons, we found that corneal afferents preferentially target parabrachial-projecting neurons in trigeminal dorsal horn. These results suggest that nociceptive pain from the cornea may be primarily mediated by a non-thalamic ascending pathway.

Trigeminal nucleus caudalis anatomy: guidance for radiofrequency dorsal root entry zone lesioning.

OBJECT: This study seeks to improve the accuracy of trigeminal nucleus caudalis dorsal root entry zone (DREZ) radiofrequency lesioning by quantifying the size and orientation of the nucleus caudalis. METHODS: Using serial axial photographs of 6 formalin-fixed cadaver brainstems, digital nucleus caudalis measurements were taken at 1-mm intervals from the level of the obex to the C(2) dorsal nerve roots. RESULTS: From the obex to the C(2) dorsal nerve roots, the nucleus caudalis decreases in width (from 2.6 ± 0.2 to 1.0 ± 0.3 mm) and, excluding superficial tract thickness, decreases in axial nucleus depth (from 2.4 ± 0.3 to 1.7 ± 0.2 mm). At levels between the obex and 10 mm caudal to the obex, the accessory nerve rootlets exit the brainstem at the junction of the spinal trigeminal tract and the dorsal spinocerebellar tract. CONCLUSION: This study details the anatomic dimensions and orientation of the nucleus caudalis for surgeons who perform DREZ lesioning.

Differential localization of vesicular glutamate transporters and peptides in corneal afferents to trigeminal nucleus caudalis.

Trigeminal afferents convey nociceptive information from the corneal surface of the eye to the trigeminal subnucleus caudalis (Vc). Trigeminal afferents, like other nociceptors, are thought to use glutamate and neuropeptides as neurotransmitters. The current studies examined whether corneal afferents contain both neuropeptides and vesicular glutamate transporters. Corneal afferents to the Vc were identified by using cholera toxin B (CTb). Corneal afferents project in two clusters to the rostral and caudal borders of the Vc, regions that contain functionally distinct nociceptive neurons. Thus, corneal afferents projecting to these two regions were examined separately. Dual immunocytochemical studies combined CTb with either calcitonin gene-related peptide (CGRP), substance P (SP), vesicular glutamate transporter 1 (VGluT1), or VGluT2. Corneal afferents were more likely to contain CGRP than SP, and corneal afferents projecting to the rostral region were more likely to contain CGRP than afferents projecting caudally. Overall, corneal afferents were equally likely to contain VGluT1 or VGluT2. Together, 61% of corneal afferents contained either VGluT1 or VGluT2, suggesting that some afferents lack a VGluT. Caudal corneal afferents were more likely to contain VGluT2 than VGluT1, whereas rostral corneal afferents were more likely to contain VGluT1 than VGluT2. Triple-labeling studies combining CTb, CGRP, and VGluT2 showed that very few corneal afferents contain both CGRP and VGluT2, caudally (1%) and rostrally (2%). These results suggest that most corneal afferents contain a peptide or a VGluT, but rarely both. Our results are consistent with a growing literature suggesting that glutamatergic and peptidergic sensory afferents may be distinct populations.

Cervicogenic headache: evidence that the neck is a pain generator.

This review was developed as part of a debate, and takes the "pro" stance that abnormalities of structures in the neck can be a significant source of headache. The argument for this is developed from a review of the medical literature, and is made in 5 steps. It is clear that the cervical region contains many pain-sensitive structures, and that these are prone to injury. The anatomical and physiological mechanisms are in place to allow referral of pain to the head including frontal head regions and even the orbit in patients with pain originating from many of these neck structures. Clinical studies have shown that pain from cervical spine structures can in fact be referred to the head. Finally, clinical treatment trials involving patients with proven painful disorders of upper cervical zygapophysial joints have shown significant headache relief with treatment directed at cervical pain generators. In conclusion, painful disorders of the neck can give rise to headache, and the challenge is to identify these patients and treat them successfully.

Morphological interrelationship between astrocytes and nerve endings in the rat spinal trigeminal nucleus caudalis.

It has been reported that the spinal trigeminal nucleus caudalis (Sp5C), which receives nociceptive information from the oro-facial regions, has four laminae. To clarify the role of glial cells in the transmission of the nociceptive information, the present study was conducted to examine the detailed distribution of astrocytes in each lamina and also to investigate a morphological interrelationship between the astrocytes and nerve endings in the rat Sp5C. After the preparation of the serial cryostat sections, immunohistochemistry for glial fibrillary acidic protein (GFAP) was employed to identify the astrocytes, and immunohistochemistry for substance P (SP), calcitonin gene-related peptide (CGRP), was used for the nerve endings. We also employed double-labeling immunofluorescence and electron microscopic immunohistochemistry for the GFAP/SP or GFAP/CGRP. GFAP-positive reactions were observed in all laminae of the Sp5C, and SP- or CGRP-positive nerve endings were observed in the lamina I and II. Additionally, we clarified the presence of GFAP/SP- or GFAP/CGRP-positive reactions by the double-labeling immunofluorescence and demonstrated the morphological interrelationship between the astrocytes and nerve endings by the double-labeling electron microscopic immunohistochemistry. These findings suggest that astrocytes might play some roles in the transmission of nociceptive information from the oro-facial region.

Differential responses of rostral subnucleus caudalis and upper cervical dorsal horn neurons to mechanical and chemical stimulation of the parotid gland in rats.

Blockage of the salivary duct can produce pain and inflammation from the build up of saliva in the parotid gland. The processing of parotid inflammation-induced pain, however, is poorly understood. The purpose of this study was to clarify the functional involvement of the trigeminal subnucleus interpolaris/caudalis transition region (Vi/Vc) and upper cervical spinal cord (C1/C2) in processing nociceptive input relevant to parotitis. The effect of capsaicin-induced parotitis was examined on a total of 37 nociceptive neurons isolated from the Vi/Vc (n = 23) and C1/C2 (n = 14) regions. Eight of 23 Vi/Vc neurons responded to mechanical distention of the parotid gland, whereas no C1/C2 neurons responded to the parotid distention. Receptive field characteristics in all neurons were examined following capsaicin injections into the parotid gland. Mechanical and cold responses increased significantly in C1/C2 but not Vi/Vc neurons following capsaicin. Receptive field sizes also increased in C1/C2 but not Vi/Vc neurons. At the Vi/Vc transition region, pinch-evoked activity increased in neurons receiving convergent inputs from the parotid gland and facial skin when compared to non-convergent neurons. The present data indicate that the hyperalgesia and referred pain associated with parotitis may result from sensitization of C1/C2, but not Vi/Vc nociceptive neurons.

Postnatal development of excitation propagation in the trigeminal subnucleus caudalis evoked by afferent stimulation in mice.

The postnatal development of nociceptive afferent activity expansion and its modulation features were examined in mice using an optical imaging technique. Developing mice (1-2 weeks old (N1-2 w), 3-4 weeks old (N3-4 w), 5-6 weeks old (N5-6 w) and 7-8 weeks old (N7-8 w)) and neonatally capsaicin-treated mice were used. The propagation of neuronal excitation was measured by changes in fluorescent intensity in horizontal brain stem slices evoked by electrical stimulation to the trigeminal spinal tract. A single-pulse stimulation evoked excitation propagation in the trigeminal caudalis (Vc). The propagation area was larger in N1-2 w than in N7-8 w, and no differences were observed between capsaicin-treated and naive mice in the same age groups. Repetitive stimulation (100 Hz, 30 pulses) elicited long-lasting and widespread excitation propagation. The excitation propagation area was significantly larger in N7-8 w than in N1-2 w, N3-4 w and N5-6 w. This propagation was suppressed by 5 microM L-703.606, an NK1-receptor antagonist, suggesting that the repetitive stimulation-elicited excitation may require substance-P releases. Morphological observations demonstrated that the neural network in the Vc had grown by postnatal week 5. These results suggest that nociceptive afferent activity co-operatively matures with development of the network structure in the Vc, and that a mechanism for prolonged increase in central excitability is established during a later postnatal period.

Coeruleotrigeminal inhibition of nociceptive processing in the rat trigeminal subnucleus caudalis.

It has been accepted that the descending system from the nucleus locus coeruleus (LC)/nucleus subcoeruleus (SC) plays a significant role in spinal nociceptive processing. The present study was designed to examine modulation of nociceptive processing in the caudal part of the trigeminal sensory nuclear complex, the trigeminal subnucleus caudalis which is generally considered to be involved in the relay of oral-facial nociceptive information. Experiments were performed on anesthetized Sprague-Dawley rats. The site of LC/SC stimulation was confirmed by histology using potassium ferrocyanide to produce a Prussian blue reaction product marking the iron deposited from the stimulating electrode tip. Only data from rats which had electrode placements in the LC/SC were used. Electrical stimulation was delivered at a stimulus intensity below 100 microA in the present study. Stimulation at sites inside the LC/SC produced a reduction of both spontaneous activity and responses of subnucleus caudalis neurons to somatic input, especially nociceptive input. Increasing stimulation frequency in the LC/SC resulted in an increase in inhibitory effects on nociceptive responses of subnucleus caudalis neurons. At three of nine sites outside the LC/SC, electrical stimulation was effective on descending inhibition. A significant difference in the inhibitory effects was observed when the inhibitory effects were compared between sites of stimulation inside the LC/SC and three effective sites of stimulation outside the LC/SC. These findings suggest that nociceptive processing in the subnucleus caudalis is under the control of the descending modulation system from the LC/SC. To understand the effects of repetitive stimulation with high frequency on fine unmyelinated LC/SC fibers, the existence of recurrent collateral excitation in the LC/SC may be considered.

An immunocytochemical investigation of human trigeminal nucleus caudalis: CGRP, substance P and 5-HT1D-receptor immunoreactivities are expressed by trigeminal sensory fibres.

5-HT1D (but not 5-HT1B)-receptor immunoreactivity (i.r.) can be detected on trigeminal fibres within the spinal trigeminal tract of the human brainstem. The present study used immunohistochemical and morphometric techniques to determine the proportions of trigeminal fibres expressing substance P, CGRP or 5-HT1D-receptor immunoreactivities. Co-localization studies between 5-HT1D-receptor and substance P- or CGRP-i.r. were also performed. Brainstem material was obtained with consent (four donors) and the total number of immunoreactive fibres within the trigeminal tract was estimated using random field sampling. A greater proportion of fibres (>1 microm diameter) expressed CGRP-i.r. (80 +/- 6%) compared with substance P-i.r. (46 +/- 7%) or 5-HT1D-receptor-i.r. (25 +/- 1%). 5-HT1D-receptor-i.r. was co-localized on some CGRP- or substance P-i.r. fibres. This suggests that 5-HT1D-receptors can regulate the release of CGRP and substance P and may be relevant to the clinical effectiveness of 5-HT1B/1D-receptor agonists in the treatment of migraine and other cranial pain syndromes.

Organization of choline acetyltransferase-containing structures in the cranial nerve motor nuclei and spinal cord of the monkey.

Cholinergic structures in the cranial nerve motor nuclei and ventral and lateral horns of the spinal cord of the monkey, Macaca fuscata, were investigated immunohistochemically with a monoclonal antibody against monkey choline acetyltransferase (ChAT). ChAT-immunoreactive perikarya and dendrites were present in the oculomotor, trochlear, abducent, trigeminal motor, facial and hypoglossal nuclei, nucleus of Edinger-Westphal, nucleus ambiguus, dorsal nucleus of the vagus, lamina IX of the cervical, thoracic and lumbar spinal cords, and intermediolateral nucleus of the thoracic spinal cord. The neuropil of the trigeminal motor, facial and hypoglossal nuclei, nucleus ambiguus and lamina IX of the cervical, thoracic and lumbar spinal cords contained many ChAT-positive bouton-like structures and they were seemingly in contact with perikarya and dendrites of motoneurons, suggesting that motoneurons in these nuclei are cholinoceptive as well as cholinergic. The oculomotor, trochlear and abducent nuclei, nucleus of Edinger-Westphal, dorsal nucleus of the vagus and intermediolateral nucleus of the thoracic spinal cord contained a small number of ChAT-immunoreactive bouton-like structures, but they did not contact with perikarya and dendrites of ChAT-positive neurons. These observations suggest that the organization of the motor nuclei is complex, at least regarding the cholinoceptivity.

The ganglionic origins and central projections of primary sensory neurons innervating the upper and lower lips in the rat.

Retrograde and transganglionic transport of horseradish peroxidase (HRP) was used to investigate the neurons innervating the upper and the lower lips and their central projections in the rat. Both the upper and the lower lips were observed to be innervated by a very large number of trigeminal sensory neurons, with their cell bodies located in the maxillary and the mandibular parts of the trigeminal ganglion, respectively. The central projections of neurons innervating the upper lip formed a long continuous column starting rostrally at midlevels of the trigeminal main sensory nucleus (5P) and extending caudally through the C1 dorsal horn, with occasional fibers reaching the C3 segment. The heaviest projections appeared in the middle portions of 5P and nucleus interpolaris (5I), as well as in the rostral part of nucleus caudalis (5C). A small but consistent projection to the solitary tract nucleus, originating from cells in the inferior vagal ganglion, was observed in the upper-lip experiments. The central projections from neurons innervating the lower lip also appeared as a long column located dorsally or dorsomedially to the projections from the upper lip. The most prominent projections from the lower lip were located in the caudal part of 5P, the middle part of 5I, and the caudal two-thirds of 5C. Sparse projections could be traced as far caudally as C4. At 5C and cervical levels, some labeling appeared contralaterally in the same location as on the ipsilateral side.

The anatomical basis for cervicogenic headache.

The neuroanatomical basis for cervicogenic headache is convergence in the trigeminocervical nucleus between nociceptive afferents from the field of the trigeminal nerve and the receptive fields of the first three cervical nerves. Only structures innervated by C1-C3 have been shown to be capable of causing headache. These are the muscles, joints and ligaments of the upper three cervical segments, but also include the dura mater of the spinal cord and posterior cranial fossa and the vertebral artery.

Corneal representation within the trigeminal subnucleus caudalis and adjacent bulbar lateral reticular formation of the cat.

Corneal units in the trigeminal subnucleus caudalis and adjacent bulbar lateral reticular formation were studied in urethane-chloralose anesthetized cats. Corneal units were categorized into four classes: low-threshold corneal (LTC) units, high-threshold corneal (HTC) units, wide dynamic range (WDR) units with corneal input, and subnucleus reticularis ventralis (SRV) units with corneal input. Corneal receptive fields of these four classes of corneal afferent units consisted of 3-6 spots. Mechanical thresholds of LTC units were lower than 30 mg (2.6 g/mm2) and were comparable to the sensory threshold of the human cornea measured in patients with cataract. Mechanical thresholds of the other 3 classes of corneal afferent units were well above the pain threshold in the human cornea. LTC units were located in the magnocellular layer of trigeminal subnucleus caudalis and were intermingled with cutaneous low-threshold mechanoreceptive units. HTC units were coexistent with nociceptive specific units in the marginal layer and in the outer zone of substantia gelatinosa. WDR units with corneal input were found in the lateral part of trigeminal lamina V equivalent, which corresponds to the lateral part of subnucleus reticularis dorsalis. These 3 classes of corneal units were found at a level 2.7-3.5 mm caudal to the obex. SRV units were found in the dorsolateral part of SRV along the entire length of the medulla oblongata caudal to the obex. These results support the suggestion that either nonpainful sensation or pain can be evoked from the cornea.

Biochemical and anatomical consequences of adult infraorbital nerve transection for serotonergic afferents within rat trigeminal subnuclei interpolaris and caudalis.

Immunocytochemistry and high-performance liquid chromatography with electrochemical detection (HPLC-ED) were used, more than 76 days after infraorbital nerve (ION) transection, to examine the distribution and density of serotonin-immunoreactive (5-HTIR) axons, as well as serotonin (5-HT) and 5-hydroxyindoleacetic acid (5-HIAA) content, within the infraorbital (IO) regions of subnuclei caudalis (SpVc) and interpolaris (SpVi). In SpVi, increases in 5-HT concentration and in density of 5-HTIR axonal varicosities were observed on the lesioned side. No changes were seen in SpVc.

Neural pathway for aggressive display in Betta splendens: midbrain and hindbrain control of gill-cover erection behavior.

Horseradish peroxidase (HRP) was used to identify parts of the presumptive neural pathway for gill cover erection, a behavioral display pattern performed by Siamese fighting fish (Betta splendens) during aggressive interactions. Motor, motor integration and sensory areas were identified in the medulla and mesencephalon. Motor neurons of the dilator operculi muscle, the effector muscle for gill cover erection, are located in the lateral and medial parts of the caudal trigeminal motor nucleus. Iontophoretic injections of HRP into the lateral trigeminal motor nucleus resulted in labeled cell bodies in two motor areas (medial part of the trigeminal motor nucleus, anterior part of the motor nucleus of cranial nerve IX-X), two parts of the reticular formation (medial and inferior reticular areas), and two nuclei of the octavolateralis system (nucleus medialis, magnocellular octaval nucleus). The HRP injections in the medial part of the caudal trigeminal motor nucleus resulted in labeled cells in the lateral part of the nucleus and in the medial reticular nucleus. Discrete injections of HRP into nucleus medialis revealed a strong axonal projection that terminated in the torus semicircularis. The medial reticular area and both of the octavolateralis nuclei received projections from their contralateral counterparts. Connections between motor areas, and between parts of the reticular formation, may coordinate the performance of gill cover erection with other behavioral patterns used during aggressive display. Connections with the octavolateralis system may provide information on the strength of an opponent's tail beats via the lateral-line system, as well as vestibular information about the fish's own orientation during aggressive display. The organization of inputs to the trigeminal motor nucleus in Betta, a perciform fish, was found to differ from that reported in the common carp, a cypriniform fish. These differences may underlie the different behavioral capabilities of the two groups of fish.

Intersubnuclear connections within the rat trigeminal brainstem complex.

Prior intracellular recording and labeling experiments have documented local-circuit and projection neurons in the spinal trigeminal (V) nucleus with axons that arborize in more rostral and caudal spinal trigeminal subnuclei and nucleus principalis. Anterograde tracing studies were therefore carried out to assess the origin, extent, distribution, and morphology of such intersubnuclear axons in the rat trigeminal brainstem nuclear complex (TBNC). Phaseolus vulgaris leucoagglutinin (PHA-L) was used as the anterograde marker because of its high sensitivity and the morphological detail provided. Injections restricted to TBNC subnucleus caudalis resulted in dense terminal labeling in each of the more rostral ipsilateral subnuclei. Subnucleus interpolaris projected ipsilaterally and heavily to magnocellular portions of subnucleus caudalis, as well as subnucleus oralis and nucleus principalis. Nucleus principalis, on the other hand, had only a sparse projection to each of the caudal ipsilateral subnuclei. Intersubnuclear axons most frequently traveled in the deep bundles within the TBNC, the V spinal tract, and the reticular formation. They gave rise to a number of circumscribed, highly branched arbors with many boutons of the terminal and en passant types. Retrograde single- or multiple-labeling experiments assessed the cells giving rise to TBNC intersubnuclear collaterals. Horseradish peroxidase (HRP) and/or fluorescent tracer injections into the thalamus, colliculus, cerebellum, nucleus principalis, and/or subnucleus caudalis revealed large numbers of neurons in subnuclei caudalis, interpolaris, and oralis projecting to the region of nucleus principalis. Cells projecting to more caudal spinal trigeminal regions were most numerous in subnuclei interpolaris and oralis. Some cells in lamina V of subnucleus caudalis and in subnuclei interpolaris and oralis projected to thalamus and/or colliculus, as well as other TBNC subnuclei. Such collateral projections were rare in nucleus principalis and more superficial laminae of subnucleus caudalis. TBNC cells labeled by cerebellar injections were not double-labeled by tracer injections into the thalamus, colliculus, or TBNC. These findings lend generality to currently available data obtained with intracellular recording and HRP labeling methods, and suggest that most intersubnuclear axons originate in TBNC local-circuit neurons, though some originate in cells that project to midbrain and/or diencephalon.

Polysensory properties of neurons in the anterior bank of the caudal superior temporal sulcus of the macaque monkey.

1. We examined the sensory properties of cells in the anterior bank of the caudal part of the superior temporal sulcus (caudal STS) in anesthetized, paralyzed monkeys to visual, auditory, and somesthetic stimuli. 2. In the anterior bank of the caudal STS, there were three regions distinguishable from each other and also from the middle temporal area (MT) in the floor of the STS and area Tpt in the superior temporal gyrus. The three regions were located approximately in the respective inner, middle, and outer thirds of the anterior bank of the caudal STS. These three regions are referred to, from the inner to the outer, as the medial superior temporal region (MST), the mostly unresponsive region, and the caudal STS polysensory region (cSTP), respectively. 3. The extent of MST and its response properties agreed with previous studies. Cells in MST responded exclusively to visual stimuli, had large visual receptive fields (RFs), and nearly all (91%) showed directional selectivity. 4. In the mostly unresponsive region, three quarters of cells were unresponsive to any stimulus used in this study. A quarter of the cells responded to only visual stimuli and most did not show directional selectivity for moving stimuli. Several directionally selective cells responded to movements of three-dimensional objects, but not of projected stimuli. 5. The response properties of cells in the superficial cortex of the caudal superior temporal gyrus, a part of area Tpt, external to cSTP were different from those of cells in the three regions in the anterior bank of the STS. Cells in Tpt were exclusively auditory, and had much larger auditory RFs (mean = 271 degrees) than those of acoustically-driven cSTP cells (mean = 138 degrees). 6. The cSTP contained unimodal visual, auditory, and somesthetic cells as well as multimodal cells of two or all three modalities. The sensory properties of cSTP cells were as follows. 1) Out of 200 cells recorded, 102 (51%) cells were unimodal (59 visual, 33 auditory, and 10 somesthetic), 36 (18%) cells were bimodal (21 visual+auditory, 7 visual+somesthetic, and 8 auditory+somesthetic), and four (2%) cells were trimodal. Visual and auditory responses were more frequent than somesthetic responses: the ratio of the population of cells driven by visual: auditory: somesthetic stimuli was 3:2:1. 2) Visual RFs were large (mean diameter, 59 degrees), but two-thirds were limited to the contralateral visual hemifield. About half the cells showed directional selectivity for moving visual stimuli. None showed selectivity for particular visual shapes.(ABSTRACT TRUNCATED AT 400 WORDS)

Direct projections from the caudal spinal trigeminal nucleus to the striatum in the cat.

The results of a WGA-HRP and HRP study in the cat indicated that some neurons in the marginal zone (lamina I) of the caudal spinal trigeminal nucleus sent their axons contralaterally to the striatum; mainly to the dorsal part of the putamen, and additionally to the ventrolateral part of the caudate nucleus, at the stereotaxic rostrocaudal levels of A 13.0-A 15.5.