Properties of nociceptive and non-nociceptive neurons in trigeminal subnucleus oralis of the rat

Recent studies have provided evidence suggesting the involvement of rostral components of the V brainstem complex such as trigeminal (V) subnucleus oralis in orofacial pain mechanisms. Since there has been no detailed investigation of the possible existence of nociceptive oralis neurons in the rat to substantiate this recent evidence, the present study was initiated to determine if neurons responsive to noxious orofacial stimuli were present in subnucleus oralis and to characterize their functional properties. In anesthetized rats, recordings were made of the extracellular activity of single neurons functionally characterized as low-threshold mechanoreceptive (LTM), wide dynamic range (WDR) or nociceptive-specific (NS) neurons. The 342 LTM neurons responded only to light mechanical stimulation of orofacial tissues. The mechanoreceptive field of the LTM neurons included the intraoral region in 28% and was localized to the adjacent perioral area in 65%. For 95% the field was localized within one V division. Responses evoked in LTM neurons by electrical stimulation of the orofacial mechanoreceptive field revealed A fiber afferent inputs but no activity that could be attributed to C fiber afferent inputs. The 72 nociceptive neurons included 52 WDR neurons which responded to light (e.g. tactile) as well as noxious (e.g. heavy pressure; pinch) mechanical stimulation of perioral cutaneous and intraoral structures, and 20 NS neurons which responded exclusively to noxious mechanical stimuli. They also differed from the LTM neurons in that 36% of the WDR and 20% of the NS neurons had a mechanoreceptive field involving more than one V division. However, in accordance with our findings for the LTM neurons, the majority of WDR and NS neurons had a mechanoreceptive field involving the intraoral and perioral representations of the mandibular and/or maxillary divisions; those neurons having a mandibular field which especially included intraoral structures predominated in the dorsomedial zone of subnucleus oralis whereas those with a perioral mechanoreceptive field which particularly involved the maxillary division were concentrated in the ventrolateral zone of oralis. In contrast to the LTM neurons, 57% of the WDR and 67% of the NS neurons showed evidence of electrically evoked C fiber as well as A fiber afferent inputs from their mechanoreceptive field. We also noted suppression of the electrically evoked responses by heating of the tail or pinching of the paw. This effect was considered to be a reflection of diffuse noxious inhibitory controls, and was seen in NS as well as WDR neurons; most, but not all, of these neurons received A fiber as well as C fiber orofacial afferent inputs.(ABSTRACT TRUNCATED AT 400 WORDS)     

Laminar distribution of receptive field properties in the primary visual cortex of the mouse

We studied the receptive field properties of single neurons in the primary visual cortex (area 17) of the mouse and the distribution of receptive field types among the cortical laminae. Three basic receptive field types were found: 1) Cells with oriented receptive fields, many of which could be classified as simple or complex, were found in all layers of the cortex, but occurred with greater frequency in layers II and III and less commonly in Layer IV. 2) Cells with non-oriented receptive fields had ON, OFF, or ON-OFF centers; they were found in all layers but were predominant in layer IV. Two subclasses of non-oriented receptive fields were characterized based on their responses to stationary and moving stimuli. One group of cells with non-oriented receptive fields responded vigorously with sustained firing to stationary flashing stimuli, and also responded well to moving stimuli over a wide range of stimulus velocities. A second group of non-oriented cells, termed motion-selective, responded poorly or not at all to stationary stimuli and responded optimally to moving stimuli over a restricted range of velocities. 3) A distinct group of neurons, termed large field, non-oriented (LFNO) cells, were found almost exclusively in layer V. LFNO cells had receptive fields that were larger than those of the other two major classes at all visual-field locations; they also had higher rates of spontaneous activity and responded to higher stimulus velocities than the other classes. In these respects, LFNO cells resembled the layer V cells of area 17 in the cat and the layer V and VI cells of area 17 in the monkey that project to the superior colliculus. We injected horseradish peroxidase into the superior colliculus, and determined that corticotectal cells in the mouse were also located in layer V, the layer where we recorded LFNO cells. Additional evidence that some LFNO cells project to the superior colliculus was provided by preliminary experiments in which we stimulated the superior colliculus and antidromically activated cortical cells with LFNO receptive fields. Neurons with LFNO receptive fields thus constitute a class that is functionally distinct, with cell bodies that are located in a single layer (V) of area 17 in the mouse.     

Superior colliculus activity related to attention and to connotative stimulus meaning

Single- and multi-unit activity was recorded from cells in the superior colliculus of two awake monkeys (Macaca fascicularis). 32.5% of 366 investigated units responded while the animals attentively gazed at visual stimuli. 50% of these neurons responded to all stimuli presented, including stationary and moving light bars, whereas the other neurons only responded to specific stimuli like faces or food. The responses of a part of these neurons depended on the connotative stimulus meaning.     

Laminar distribution and the response of superior colliculus neurons to stimulation of the periodontal ligament in the rabbit

Unit discharges of superior colliculus neurons fired by electrical stimulation of the periodontal ligament and/or mechanical stimulation of the mandibular incisor tooth were extracellularly recorded, and their laminar distribution and response properties were examined. The neurons were widely distributed in the deep layer of the anterior two-thirds of the superior colliculus. Most of them were located in the lateral site of the profound layer, a few in the lateral site of the intermediate layer. Their locations correspond to the lower nasal quadrant of the visual field. The mean first-spike latency of the intermediate layer neurons (7.2 ms) elicited by electrical stimulation of the periodontal ligament was shorter than that of the profound-layer neurons (13.8 ms). About one-third of the neurons were suppressed by conditioning electrical stimulation of the optic chiasma. These results suggested that in addition to visual, auditory, and body surface tactile sensations, intraoral somesthetic sensations also participate in the functioning of the superior colliculus which appears to play a role in orienting the animal toward visual, somesthetic, and auditory stimuli.     

Connections of the superior olivary complex in the rufous horseshoe bat Rhinolophus rouxi

In the rufous horseshoe bat (Rhinolophus rouxi), the superior olivary complex contains four main divisions. In comparison with other species, the most lateral division is clearly homologous to the lateral superior olive (LSO); the most medial division is homologous to the medial nucleus of the trapezoid body (MNTB). Lying between these landmarks, in approximately the position of the medial superior olive (MSO) of other mammals, are two additional divisions that are cytoarchitecturally distinct from one another yet do not greatly resemble the MSO of nonecholocating mammals such as the cat. We refer to these nuclei as the dorsal medial superior olive (DMSO) and the ventral medial superior olive (VMSO). We examined the afferent and efferent connections of all of these cell groups with retrograde and anterograde transport of WGA-HRP from the superior olivary complex. In the same animals we recorded the binaural response properties of single units in the superior olivary complex. Virtually all units recorded in LSO were excitatory to the ipsilateral ear and inhibitory to the contralateral ear (EI); all of the units sampled in the MNTB and most of those sampled in the VMSO responded only to the contralateral ear (OE). In DMSO the binaural properties of units were varied: the number of units that were inhibitory to the ipsilateral ear and excitatory to the contralateral ear (IE) was about equal to the number of units excitatory to both ears (EE); a few units had OE responses; no units had EI responses. Connectional correlates for these binaural response properties are seen in the patterns of retrograde transport from WGA-HRP injections in the divisions of the superior olive. The LSO receives projections from the ipsilateral cochlear nucleus and MNTB; MNTB receives projections from the contralateral cochlear nucleus. The DMSO and VMSO both receive bilateral projections from the cochlear nuclei. The results of retrograde and anterograde transport suggest that VMSO, in addition, receives projections from the ipsilateral MNTB. The LSO, DMSO, and VMSO all project to the ventral two-thirds of the central nucleus of the inferior colliculus, and their targets are approximately coextensive. However, the LSO projects bilaterally to the inferior colliculus, whereas the medial cell groups project mainly ipsilaterally.     

Functional organization of the superior vestibular nucleus of the squirrel monkey.

The response to angular acceleration of units in the superior vestibular nucleus (SVN) of barbiturate-anesthetized, cerebellectomized squirrel monkeys was used to study the distribution of semicircualr-canal inputs to the nucleus. Some so-called intact animals had 6 active semicircular canals. In other animals, the 3 canals on one side were rendered nonresponsive by plugging. In plugged animals, superior, posterior, and horizontal-canal units were encountered on both the plugged and unplugged sides, showing that all 6 canals influence the nucleus. Most units responded bilaterally to labyrinthine polarization; 92.5% of units in intact animals responded to angular acceleration, and this incidence was not decreased in plugged animals. These results suggest that most units in the superior nucleus receive bilateral canal inputs. Convergence of influences arising in orthogonally related canals was detected in less than 10% of units, so the bilateral ampullary influences must arise in parallel canals. Most SVN canal units on the plugged and unplugged sides gave type I responses, indicating that the contralateral canal influence is carried by a crossed inhibitory pathway. Most units influenced by the ipsilateral superior canal were located in the lateral half of the SVN. Posterior-canal units were in the medial half. There was no clear localization of the relatively few horizontal-canal units which were encountered.     

The rostral part of the trigeminal sensory complex is involved in orofacial nociception

Single units responsive to noxious mechanical stimulation of orofacial receptive fields were recorded within the ventrobasal complex of the rat thalamus. The induced activities were compared before and after deafferentation of the subnucleus caudalis by a trigeminal tractotomy performed at the obex level. The receptive fields activated by noxious stimulation were classified as 'oral' when included in the oral, perioral or paranasal areas, and as 'facial' when included in facial regions distant from the oral cavity. After tractotomy, the unit responses to noxious stimulation of an oral field remained unchanged in 8 cases, decreased in 3 cases, and were suppressed in 4 cases. For units responding to noxious stimulation of a facial field, the responses were suppressed in 8 cases, decreased in two cases and remained unchanged in two other cases. So it appears that the rostral part of the trigeminal sensory complex (1) receives nociceptive afferents mainly from the oral and perioral areas and (2) is a relay in ascending pathways which convey painful sensations.     

Nociceptive responses of trigeminal neurons in SII-7b cortex of awake monkeys

A cluster of trigeminal nociceptive neurons was located in the lateral sulcus on the upper bank of the frontoparietal operculum in a region bordering between cortical areas SII and 7b. These neurons were isolated in cortical cell layers IV and V-VI. All nociceptive neurons responded exclusively to noxious mechanical stimulation of cutaneous receptive fields on the face/head or intraoral tissue. Sustained noxious mechanical stimulation elicited slowly adapting responses that accurately encoded the duration of the stimulation. Prolonged discharges following removal of noxious stimulation were not observed. These nociceptive specific neurons poorly encoded graded noxious stimuli. Trigeminal somatosensory neurons within and surrounding the SII-7b cluster were not topographically organized according to divisions of the trigeminal nerve, laterality of receptive fields, or division of face/head and intraoral receptive fields. The thalamocortical, corticocortical and indirect corticolimbic connectivities of SII and area 7b and the possible role of SII-7b nociceptive neurons in learning, memory and avoidance behaviors are discussed.     

Functional influences of the visuocortical projection on superior colliculus neurons in the rabbit

The effects of electrical stimulation of the visual cortex on superior colliculus neurons were investigated in adult Dutch-belted rabbits. Single units were recorded in the superior colliculus and classified as to receptive field type. Stimulation of the ipsilateral visual cortex activated 29% of the recorded superior colliculus units. No units were driven by stimulation of the contralateral visual cortex. Comparison of the relative proportional distributions of cortically driven and not driven cells having various receptive field types revealed an over-representation of driven motion type cells. The excitatory influence of the visuocortical projection to the superior colliculus in the rabbit shows a preference for neurons responsive to moving visual stimuli.     

Alterations in receptive field properties of superior colliculus cells produced by visual cortex ablation in infant and adult cats

To determine if functional alterations in the superior colliculus might account for recovery of visual behaviors following visual cortex removal in infant cats, the receptive field characteristics of single units in the superior colliculus of cats whose visual cortex was removed within the first week of life were compared with those of cats which sustained visual cortex lesions in adulthood and with those of normal cats. In the normal superior colliculus, 90% of all cells responded to moving stimuli irrespective of shape or orientation. Sixty-four percent of these units were directionally selective, responding well to movement in one direction but poorly or not at all to movement in the opposite direction. Ninety percent of units were binocular, the vast majority of these responding equally to stimulation of either eye or showing only slight preference for stimulation of the contralateral eye. Responses to stationary flashes of light were observed in only 33% of all visually activated cells in the normal superior colliculus. After visual cortex ablation in adult cats, only six percent of movement sensitive cells were directionally selective. Binocular preference was shifted following adult visual cortex lesions such that sixty percent of all cells responded exclusively or predominantly to stimulation of the contralateral eye. Seventy-one percent of all visually responsive units responded to stationary lights flashed on or off within their receptive field boundaries. Lesions limited primarily to area 17 had the same effect as larger lesions of visual cortex. Infant visual cortex lesions resulted in receptive field alterations similar to those observed after adult ablation. Only fifteen percent of motion sensitive units were directionally selective. Seventy-one percent responded exclusively or predominantly to stimulation of the contralateral eye. Seventy-six percent of visually responsive cells were activated by stationary light. Lesions largely confined to area 17 produced the same alterations as more extensive lesions of visual cortex. Thus, no evidence was found that the superior colliculus is involved in the functional reorganization presumed to occur following visual cortex ablation in infant cats. Recovery of visual behaviors following neonatal injury may therefore not involve alterations in the receptive fields of single cells.     

Coding for auditory space in the superior colliculus of the rat

Although the rat is often used to determine behavioural sound-localization capabilities or neuronal computation of binaural information, the representation of auditory space in the rat brain has not been investigated so far. We obtained extracellular recordings from auditory neurons in the superior colliculus of anaesthetized rats and examined them for spatial tuning characteristics and topographical order. Many neurons (73%) showed significant tuning, with a single peak in the azimuth response profiles based on spike rates and response latencies. Best azimuth values from neurons in one SC were generally tuned to contralateral and rarely to frontal or ipsilateral directions. Tuning width was mostly broad; at supra-threshold sound pressure levels (35 dB SPL), 55% of the units had a tuning width of > 120 degrees in contralateral space. Additionally, tuning width increased with stimulation intensity. A significant but considerably scattered topographical order of best azimuth directions was observed in the deep layers of the superior colliculus with frontal directions being represented closer to the rostral pole. Tuned auditory units in the intermediate layers of the superior colliculus, however, showed no systematic spatial arrangement. This pattern was confirmed by analysing best azimuth directions from simultaneously recorded units. Our results indicate that the rat superior colliculus contains a representation of auditory space which is similar to that described for other small mammals.     

The effect of hemidecortication on the inhibitory interactions in the superior colliculus of the cat.

Single neurons were recorded extracellularly from the superficial layers of the superior colliculus (SC) in 21 curarized cats. Four animals were normal unoperated cats, 17 were animals in which all cortical visual areas were ablated on one side from 7 to 69 days before the electrophysiological experiments. After cortical ablation all animals were blind in the visual field contralateral to the ablated side. In both normal and hemianopsic cats the effect of a visual stimulus located very far from the excitatory part of the unit receptive field, on the neuron responses to visual stimuli was studied. The remote stimulus (extra-field stimulus) was a hand moved black spot 10 degrees in diameter. In normal animals the introduction of the extra-field stimulus in the hemifield contralateral or ipsilateral to the recorded SC produced a marked reduction of unit responses to visual stimuli presented in their receptive field. This effect was particularly strong when the extra-field stimuli were introduced in the hemifield contralateral to the recorded side. In the hemianopsic animals the neurons of the SC ipsilateral to the lesion (receptive fields in the behaviorally blind hemifield) responded well to visual stimuli, but were only weakly inhibited by the extra-field stimuli presented in the blind hemifield. The neurons of this colliculus with the exception of those in the upper part of stratum griseum superficiale were normally inhibited by stimuli presented in the normal hemifield. The neurons of the SC contralateral to the lesion responded well to visual stimuli and were normally inhibited by stimuli presented in the normal hemifield; they were virtually not affected by stimuli presented in the blind hemifield. Mechanisms responsible for the abnormal inhibitory interactions between and within colliculi after cortical lesions and the possible behavioral implications of the findings are discussed.     

Neurons in the Primate Superior Colliculus are Active Before and During Arm Movements to Visual Targets

The activity of single neurons in the superior colliculus was recorded while a rhesus monkey made arm movements to visual targets located on a screen in front of him. It was found that the activity of a subpopulation of cells was clearly related to these arm movements. The neurons began to discharge either with the onset of the movement, during the movement period, or well before the onset of electromyogram (EMG) activity and movement, and could be active for the entire duration of EMG activity. While the discharge pattern of some of these'reach'neurons was not different for movements to different target positions, other cells showed graded changes in activity depending on the direction of movement. The peak discharge rate could rise to > 100 impulses/s. Some units received somatosensory input; other reach cells exhibited a visual response and/or presaccadic activity. It is likely that the primate superior colliculus is not only involved in the initiation and control of orientating movements of the eyes but also in reaching movements of the arms.     

Latencies of visually guided saccades in unilateral hemispheric cerebral lesions

Latencies of lateral visually guided saccades were studied in 60 patients without hemianopia who had unilateral focal lesions clearly visible on computed tomographic (CT) scan that were variously located in both cerebral hemispheres. Significantly asymmetrical latencies were found in 29 patients whose lesions had damaged the deep and posterior frontal region near the corpus callosum and/or, just inferior to this region, the anterior part of the internal capsule. In the 31 other patients, including those with lesions of the frontal eye fields (FEF), latencies were not significantly asymmetrical and the lesions spared the entire region just described. These topographical features suggest that the asymmetry of latencies is due to damage in a certain portion of the efferent pathways descending from the FEF. A significant increase in bilateral latency was observed in most patients whose lesions had damaged the posterior part of the parietal cortex and/or the underlying white matter. The parietal lobe could therefore exert an excitatory bilateral action on the triggering of visually guided saccades, probably mediated via the superior colliculus. A significant decrease in the bilateral or ipsilateral latency was often found in patients whose lesions had damaged the FEF or the underlying white matter. The frontal lobe could therefore exert a predominantly inhibitory bilateral action on this triggering, probably also mediated via the superior colliculus. However, an increase in contralateral latency in some patients with subcortical frontal lesions indicates that the FEF also probably have an excitatory action. This action could be transmitted directly (or indirectly via the superior colliculus) to the reticular premotor structures by tracts decussating partly through the corpus callosum.     

Retinal X-afferents bifurcate to lateral geniculate X-cells and to the pretectum or superior colliculus in cats

The question of whether retinal X-type ganglion cell axons project via axonal bifurcation to both the dorsal lateral geniculate nucleus (LGN) and the pretectum (PT) or the superior colliculus (SC) in the cat, was studied by examining the effects of PT and/or SC stimulation on the LGN cells. X-cells that responded monosynaptically to PT or SC stimulation were encountered as follows: 29%, 26%, and 4% of the tested X-cells responded to stimulation of PT, SC, and both, respectively. For the X-cells activated from the PT or SC, the latency tended to be a little longer than optic chiasm latency. The receptive field centers of the X-cells were located within the receptive fields of the multiple units from the SC whose stimulation could activate the corresponding X-cells. The present results demonstrate that a substantial proportion of the X-type LGN cells receive excitatory inputs from the retinal X-type ganglion cell axons that branch to the PT or the SC.     

Responses of neurons in nucleus ventralis posterolateralis of the cat thalamus' to hypogastric inputs

Recordings were made from 68 units in the nucleus ventralis posterolateralis (VPL) of the cat thalamus, which responded to stimulation of hypogastric afferents. These units also received nociceptive inputs from the contralateral integument. Units which responded exclusively to hypogastric afferent inputs were not found. Thirty seven of the units were nociceptive specific (NS), and the remaining 31 were wide dynamic range (WDR) units. All of these units were located in the shell region of the lateral subdivision of the caudal VPL. NS units responding to hypogastric afferent inputs had a circumscribed cutaneous receptive field on the contralateral abdomen, gluteal region, tail or hind limb. These areas corresponded to tactile dermatomes T13-S2. Similarly, the cutaneous receptive fields of WDR units receiving hypogastric afferent inputs were distributed in the contralateral abdomen, gluteal region, tail and hind limb, with the sole exception of one unit, whose receptive field also included a part of the lower thorax. These findings extend the previous findings that the shell region of the caudal VPL of the cat thalamus constitutes a thalamic link in a visceral pain pathway, and that the visceral and cutaneous pathways share a common projection locus in the VPL.     

Differential effects of trigeminal tractotomy on Adelta- and C-fiber-mediated nociceptive responses

In this study we have tested in the rat, whether trigeminal tractotomy, which deprives the spinal trigeminal nucleus caudalis (Sp5C) of its trigeminal inputs, affected differentially nociceptive responses mediated by C- vs. Adelta-nociceptors from oral and perioral regions. Tractotomy had no effect on the threshold of the jaw opening reflex, induced by incisive pulp stimulation (Adelta-fiber-mediated), but blocked the formalin response (mainly C-fiber-mediated). These results suggest that nociceptive responses mediated by trigeminal C-fibers completely depend on the integrity of the Sp5C, while intraoral sensations triggered Adelta-fibers (especially of dental origin) are primarily processed in the rostral part of the spinal trigeminal nucleus.     

Role of the superior colliculus in analyses of space: superficial and intermediate layer contributions to visual orienting, auditory orienting, and visuospatial discriminations during unilateral and bilateral deactivations.

The superior colliculus (SC) has been implicated in spatial analyses of the environment, although few behavioral studies have explicitly tested this role. To test its imputed role in spatial analyses, we used a battery of four spatial tasks combined with unilateral and bilateral cooling deactivation of the upper and intermediate layers of the superior colliculus. We tested the abilities of cats to orient to three different stimuli: (1) moving visual, (2) stationary visual, (3) stationary white-noise aural. Furthermore, we tested the ability of the cats to discriminate the relative spatial position of a landmark. Unilateral cooling deactivation of the superficial layers of the SC induced a profound neglect of both moving and stationary visual stimuli presented in, and landmark objects located within, the contralateral hemifield. However, responses to auditory stimuli were unimpaired. Unilateral cooling deactivation of both the superficial and intermediate layers induced a profound contralateral neglect of the auditory stimulus. Additional and equivalent deactivation of the opposite SC largely restored orienting to either moving visual or auditory stimuli, and restored landmark position reporting to normal levels. However, during bilateral SC deactivation, orienting to the static visual stimulus was abolished throughout the entire visual field. Overall, unilateral SC deactivation results show that the upper and intermediate layers of the SC contribute in different ways to guiding behavioral responses to visual and auditory stimuli cues. Finally, bilateral superior colliculus deactivations reveal that other structures are sufficient to support spatial analyses and guide visual behaviors in the absence of neural operations in the superior colliculus, but only under certain circumstances.     

Anatomical distribution and sensory properties of brain stem and posterior diencephalic neurons responding to genital, somatosensory, and nociceptive stimulation in the squirrel monkey.

In chloralose-urethane-anesthetized female squirrel monkeys, 325 single units sampled from a region extending from the caudal medulla to the posterior diencephalon were examined for responsiveness to genital, rectal, innocuous somatosensory, and various forms of nociceptive stimulation. The units were highly responsive, with 84% responding to at least one stimulus type. The responsive units were widely distributed in the brain stem tegmentum, deep tectum, and posterior diencephalon. Very few neurons responded to only one type of stimulation. The patterns of convergent responsiveness to the various stimulus types were not, however, a simple random function of unit responsiveness to each type of stimulus per se. Unit responses to vaginal stimulation consisted of simple increases or decreases in firing which outlasted the duration of the probing stimulus in most cases. Some units responded more strongly to cervical than to vaginal tract stimuli. The somatic receptive fields of units responding to touch-pressure stimuli were typically bilateral and quite extensive. A forceps pinch of nociceptive intensity elicited a response from 64% of the cells, and of these, 11% showed significant linear correlations between their firing rates and increasing pinch pressure in the nociceptive intensity range. Brief, localized nociceptive thermal stimuli and needle pricks failed to elicit responses from the neurons tested. Based on a comparison between the response properties of monkey brain stem neurons and the previous findings for rat and cat neurons, it was concluded that brain stem cells display species-typical sensory characteristics which have parallels in the properties of behavioral responses of these three species to genital and other sensory stimuli. Properties of unit responses to nociceptive stimuli have implications in relation to the neural mechanisms of first and second pain.     

Differential effects of trigeminal tractotomy on Aδ- and C-fiber-mediated nociceptive responses

In this study we have tested in the rat, whether trigeminal tractotomy, which deprives the spinal trigeminal nucleus caudalis (Sp5C) of its trigeminal inputs, affected differentially nociceptive responses mediated by C- vs. Aδ-nociceptors from oral and perioral regions. Tractotomy had no effect on the threshold of the jaw opening reflex, induced by incisive pulp stimulation (Aδ-fiber-mediated), but blocked the formalin response (mainly C-fiber-mediated). These results suggest that nociceptive responses mediated by trigeminal C-fibers completely depend on the integrity of the Sp5C, while intraoral sensations triggered Aδ-fibers (especially of dental origin) are primarily processed in the rostral part of the spinal trigeminal nucleus.