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Retrograde tracing with true blue (TB) and diamidino yellow (DY) and anterograde tracing with either wheatgerm agglutinin-conjugated horseradish peroxidase (WGA-HRP) or Phaseolus vulgaris leucoagglutinin (PHA-L) were employed to investigate the projections from trigeminal nucleus principalis (PrV) and trigeminal subnucleus interpolaris (SpI) to their targets in the medial ventral posterior (VPM) and posterior (POm) nuclei of the thalamus. Many more cells in both PrV and SpI were labeled by tracer injections into VPM than into POm. Only a very small number of double-labeled neurons were observed in either PrV or SpI. However, a significantly higher percentage of SpI cells projected to POm or to both POm and VPM than was the case for PrV. Anterograde tracing with WGA-HRP showed that the projections from both PrV and SpI to VPM were much denser than those from the same nuclei to POm. Small injections of PHA-L into either PrV or SpI produced a focus of fairly dense labeling in VPM and much more diffuse terminal labeling in POm. These anatomical data provide evidence for two separate trigeminothalamic pathways, one originating from PrV and the second originating from SpI. Both of these pathways converge and diverge at the thalamic level. That is, information from the PrV pathway and from the SpI pathway are both provided to VPM in a morphologically restricted fashion and to POm in a morphologically widespread fashion.

Spinal trigeminal nucleus pars oralis (SpoV) is anatomically linked to brain circuitry thought to subserve unconditioned and conditioned nictitating membrane responses in rabbit. Single-unit recording from SpoV and adjacent reticular formation obtained during conditioning from awake, behaving animals revealed modulation of unit firing related to CS, US, and CR occurrence. SpoV participates directly in the unconditioned response and probably relays US information to other brain areas subserving conditioning. The presence of CR-related activity suggests that SpoV may participate in the CR motor output pathway, and may also provide CR-related information to cerebellum. Sensory convergence and CR-related activity in reticular formation mark this structure as a candidate locus of primary neuronal plasticity in this example of conditioning.

The cell bodies and central projections of neurons innervating the vibrissae follicles and adjacent skin in the rat were investigated by retrograde and transganglionic transport of HRP. The cell bodies of neurons innervating the vibrissa follicle via the deep vibrissa nerve (DVN) were the largest, followed by those innervating the follicle via the superficial vibrissa nerve (SVN). The smallest cell bodies were those innervating the intervibrissal skin. The DVN neurons terminated centrally as an almost uninterrupted column through the trigeminal sensory nuclear complex. The DVN projections to nucleus caudalis and C1 dorsal horn were entirely restricted to laminae III, IV, and V. Besides the projections to lamina V, the DVN projections were strictly localized somatotopically at all levels replicating the peripheral organization of the vibrissae. The SVNs projected sparsely to midlevels of the main sensory nucleus but not to nuclei oralis and interpolaris. The main SVN projections appeared in laminae I-III of nucleus caudalis. In addition, a small projection to lamina V was observed. The projections to laminae II and III were organized mediolaterally in a similar way as the DVN projections; those to laminae I and V were less restricted. The intervibrissal skin neurons projected sparsely to the caudal main sensory nucleus and to the border between nuclei oralis and interpolaris. The projections to nucleus caudalis were restricted to laminae I-III and V and were organized in a similar way as the SVN projections.

The central axon of a primary afferent neuron that responded to indentation of the glabrous skin of the lower lip in a slowly adapting fashion was intra-axonally injected with horseradish peroxidase. The labeled terminal within the subnucleus oralis was examined electron microscopically. The labeled ending had a pale axoplasm and contained clear spherical synaptic vesicles. The labeled ending formed a synaptic triad with a dendrite and an unlabeled axonal ending with pleomorphic vesicles (a mixture of oval, flattened and dense core vesicles). The labeled primary ending was presynaptic only to the dendrite, while the unlabeled ending was presynaptic to both the dendrite and the labeled primary ending.

Projections of the parvocellular reticular nucleus (PCRt) to the contralateral mesencephalic trigeminal nucleus (Me5) were studied in the rat with neurophysiological and neuroanatomical techniques. Three types of responses (classified by latencies) were recorded extracellularly in the Me5 area after electrical stimulation of the PCRt: (1) R1 fast unitary reactions (latency 0.2-0.5 ms) found throughout the entire caudal Me5 area; (2) R2 slower unitary reactions (latency 0.7-1.2 ms) recorded ventral and lateral to the caudal Me5; and (3) R3 compound potentials (latency 1.0-2.5 ms) recorded within the ventrocaudal Me5. Relayed or synaptic fields were not observed. Intracellular recordings of identified cell bodies of periodontal afferents, muscle spindle afferents and interneurones revealed no short-term postsynaptic potentials following PCRt stimulation. In some experiments jaw muscle spindle afferent activity was increased by PCRt stimulation and jaw-opening and jaw-closing reflexes were facilitated in the contralateral musculature. Neuroanatomical experiments, using Phaseolus vulgaris leucoagglutinin as an anterograde tracer, showed direct projections with intensive collateralization of PCRt fibres into the Me5 area. Synaptic contacts of PCRt fibres with primary afferent cell bodies were observed in the ventrocaudal parts of the Me5 only. The electrophysiological results are discussed in relation to the neuroanatomical findings.

Two separate studies were done on two sets of rats (33 rats, 101 cells in Expt. 1; 16 rats, 25 cells in Expt. 2). In the first study trigeminal nuclear neurons were categorized by adequate stimulus and receptive field. A total of 43.6% responded to light touch and noxious (pinch and heat) stimulation; 27.7% responded only to noxious pinch and/or heat; 8.9% responded exclusively to innocuous touch, and the remainder were non-responsive. Ninety percent of the responsive cells showed increased discharge rates in response to noxious stimulation. In the second study, after characterization of response properties, single nanoinjections of Met-enkephalin (ME) were separately made into midbrain periaqueductal gray (PAG) and nucleus reticularis paragigantocellularis (PGC). The effects of ME on spontaneous (SD) and evoked (ED) discharge were assessed, and then in 8 cells, the effects were repeated and successfully challenged with naloxone. The major findings were: (1) ME applied either to PAG or PGC causes naloxone-reversible reduction of noci-evoked ED with no effect on SD; and (2) the responses of neurons to innocuous light touch were unaffected by ME.

Horseradish peroxidase (HRP) retrograde tracing experiments in the cat demonstrated projections from the trigeminal sensory nuclei to the mesencephalic parabrachial area (PBA), which is located ventral to the inferior colliculus and dorsal to the brachium conjunctivum and includes the nucleus cuneiformis and the most lateral part of the periaqueductal gray. After HRP injection into the mesencephalic PBA, HRP-labeled neuronal cell bodies in the trigeminal sensory nuclei were mainly distributed in the interpolar and caudal spinal trigeminal nuclei (Vi and Vc) bilaterally with a clear-cut contralateral dominance. In Vi, HRP-labeled neurons were mainly distributed in the lateral and medial border regions. In Vc, labeled neurons were located mainly in laminae I and V.

In order to study the morphological characteristics and terminating patterns of the neurons of the trigeminal mesencephalic nucleus (Vme), 55 masseteric neurons in Vme in the rat were stained by intracellular injection of horseradish peroxidase (HRP). Labeled cells were distributed throughout the nucleus. These neurons were divided into three types: uni- or pseudounipolar (type A, n = 43), bipolar (type B, n = 5), and multipolar cells (type C, n = 7). Each type was further divided into two subtypes according to the largest diameter of the perikarya (type a greater than or equal to 30 microns, type b less than 30 microns). The central processes of type Aa neurons projected to the following three groups of target nuclei: 1) nuclei functioning as interneurons, including supratrigeminal nucleus (Vsup), intertrigeminal nucleus (Vint), juxta-trigeminal region (Vjux), and parvicellular nucleus of the pontomedullary reticular formation (PcRF); 2) motor nuclei, including the trigeminal motor nucleus (Vmo), accessory facial nucleus (NVIIacs), accessory abducens nucleus (NVIacs), and a small number of labeled axons in the oculomotor nucleus and trochlear nucleus; 3) sensory nuclei, including the dorsomedial part of the principal trigeminal sensory nucleus (Vpdm) and the dorsomedial part of subnucleus oralis of the trigeminal spinal nucleus (Vodm). Labeled processes were dense in the Vsup, Vmo, and Vpdm. The proprioceptive pathway of the fifth nerve is discussed. Direct projections from type Aa neurons of Vme to the Vpdm and dorsolateral part of the Vsup contribute to conduction of the proprioceptive information from spindles of masticatory muscle to the contralateral thalamus in the rat. Different axon morphology, distribution, terminal branch density, and terminating patterns of type Aa neurons were noted in different functional groups of the projecting nuclei, especially in the Vsup, Vmo, and Vpdm. The highest terminal branching density, the most extensive distribution, and two different types of branching patterns (claw-like and comb-like) were observed in Vsup. Selective distribution and single-beaded or "Y"-shaped terminal branches were observed in Vmo. In the Vppdm the axonal branches were sparser than in the Vsup or Vmo, and had an arrangement like the branches of a weeping willow tree. These characteristics of anatomical organization might be related to the function of each projecting nucleus.

The present study tested the hypothesis that the trigeminal (V) primary afferent projection to the contralateral dorsal horn originates in midline hairy skin. A prior study (Jacquin et al., 1990) showed that this crossed projection is heaviest to ophthalmic regions of medullary and cervical dorsal horns, and that it does not arise from V ganglion cells that innervate cornea, nasal mucosa, or cerebral dura mater. Here, retrograde double-labeling methods were used to show that many ophthalmic ganglion cells that innervate midline hairy skin via the supraorbital nerve project to the contralateral medullary and upper cervical dorsal horns. Diamidino yellow injections into the right dorsal horn labeled an average of 104 cells in the left V ganglion. Of these contralaterally projecting ganglion cells, an average of 45% were also labeled by horseradish peroxidase (HRP) injections into the left supraorbital nerve, and 25% were also labeled by HRP injections into the midline opthalmic hairy skin. However, only 2% were labeled by HRP injections restricted to left supraorbital vibrissae follicle nerves. Almost all of the double-labeled cells were located in the dorsal one-half of the V ganglion, and they did not differ in size from single-labeled cells. On the basis of these and prior data, we conclude that a high percentage of contralaterally projecting V ganglion cells originate in midline hairy skin. It is also likely that the contralaterally projecting V ganglion cells serve a low-threshold mechanoreceptive function, given the relatively large ganglion cells and axons giving rise to this pathway and their central terminations in dorsal horn laminae III-V.

The ventral posteromedial nucleus (VPM) of the monkey thalamus was investigated with combined immunocytochemical, histochemical, and connection-tracing techniques. Injections of anterogradely transported tracers were placed selectively in the caudal nucleus of the spinal trigeminal nuclear complex, and retrogradely transported horseradish peroxidase (HRP) or fluorescent dyes were placed on the surface or into the depths of defined parts of the trigeminal representation in the first somatic sensory area (SI) of the cerebral cortex. The results are correlated with those of the preceding paper (Rausell and Jones, 1991), which demonstrated the presence of 2 domains in the nucleus on the basis of different patterns of cytochrome oxidase (CO) staining and calcium-binding protein immunoreactivity. The cells of the CO-defined rod and matrix domains receive inputs from different components of the trigeminal afferent system and project to different layers of SI. The large- and medium-sized relay cells of the CO-rich rods, which are immunoreactive for parvalbumin, all project to middle layers of SI. The small relay cells of the weakly-stained CO-matrix, surrounding and intervening between the rods, are immunoreactive for 28-kDa calbindin and project to superficial layers (I and II) of SI. Anterograde tracing studies reveal that the rod domain in VPM is innervated by fibers arising in the contra- and ipsilateral principal trigeminal nucleus, while the matrix domain (and calbindin-positive domains in adjacent nuclei) are innervated by fibers arising in the caudal nucleus of the spinal trigeminal complex. These results demonstrate the modularity and parallel streaming of the functional components of the trigeminal part of the somatic sensory system and suggest that lemniscal and nonlemniscal elements of the system gain access by separate routes to different layers of the SI cortex.

The topographical distribution of motoneurons innervating the digastric muscle in the rabbit and guinea pig was re-examined by the retrograde tracing method of HRP (horseradish peroxidase). Motoneurons innervating the anterior belly of the digastric muscle of the rabbit and guinea pig constituted a longitudinal cell column in the ventromedial part of the motor nucleus of the trigeminal nerve. Motoneurons innervating the posterior belly of the digastric muscle were localized in the accessory facial nucleus. No motoneurons supplying the digastric muscle were found within the main facial nucleus.

The aim of the present study was to determine the brainstem afferents and the location of neurons giving rise to monoaminergic, cholinergic, and peptidergic inputs to the cat trigeminal motor nucleus (TMN). This was done in colchicine treated animals by using a very sensitive double immunostaining technique with unconjugated cholera-toxin B subunit (CT) as a retrograde tracer. After CT injections in the TMN, retrogradely labeled neurons were most frequently seen bilaterally in the nuclei reticularis parvicellularis and dorsalis of the medulla oblongata, the alaminar spinal trigeminal nucleus (magnocellular division), and the adjacent pontine juxtatrigeminal region and in the ipsilateral mesencephalic trigeminal nucleus. We further observed that inputs to the TMN arise from the medial medullary reticular formation (the nuclei retricularis magnocellularis and gigantocellularis), the principal bilateral sensory trigeminal nucleus, and the dorsolateral pontine tegmentum. In addition, the present study demonstrated that the TMN received 1) serotonergic afferents, mainly from the nuclei raphe obscurus, pallidus, and dorsalis; 2) catecholaminergic afferent projections originating exclusively in the dorsolateral pontine tegmentum, including the Kölliker-Fuse, parabrachialis lateralis, and locus subcoeruleus nuclei; further, that 3) methionin-enkephalin-like inputs were located principally in the medial medullary reticular formation (nuclei reticularis magnocellularis and gigantocellularis and nucleus paragigantocellularis lateralis), in the caudal raphe nuclei (Rpa and Rob) and the dorsolateral pontine tegmentum; 4) substance P-like immunoreactive neurons projecting to the TMN were present in the caudal raphe and Edinger-Westphal nuclei; and 5) cholinergic afferents originated in the whole extent of the nuclei reticularis parvicellularis and dorsalis including an area located ventral to the nucleus of the solitary tract at the level of the obex. In the light of these anatomical data, the present report discusses the possible physiological involvement of TMN inputs in the generation of the trigeminal jaw-closer muscular atonia occurring during the periods of paradoxical sleep in the cat.

1. These experiments were based on the findings that antidromic firing is observed in first-order sensory afferents during fictive locomotion and were designed to test the hypothesis that a similar central modulation of afferent discharge occurs during mastication. To do this, spindle afferents were recorded in the trigeminal mesencephalic nucleus (Mes V) of anesthetized and paralyzed rabbits during fictive mastication. The cortical masticatory area was stimulated to induce mastication, and activity of the XIIth or the Vth nerves were recorded to monitor the masticatory motor rhythm. 2. Although we could find little evidence that antidromic discharges invade the somatic region of this class of sensory afferents, we did discover a previously unrecognized type of modulation of afferent firing. 3. Of 83 slowly adapting muscle spindle afferents, 33 were modulated during fictive mastication. In 28 cases, the modulation consisted of a phasic inhibition, whereas for the remaining units it could be either a phasic excitation (n = 2) or an excitation alternating with an inhibition (n = 3). 4. Rapidly adapting units were also tested when encountered. Tonic or phasic excitation was never observed. The presence of inhibition could not be verified for this population because tonic activity could not be maintained by passive stretch. 5. The main electroneurogram (ENG) burst of the XIIth and Vth cranial nerves occurred during the opening phase of the masticatory cycle, and in all cases where the records were clear (22 out of 33), phasic inhibition of the afferents coincided with the ENG burst. 6. There was no difference in the distributions of the modulated and of the unmodulated units along the length of the Mes V nucleus. 7. Approximately 40% of trigeminal spindle afferent cell bodies have dendrites, and we suggest that these are the ones rhythmically modulated during fictive mastication. The possible role of this modulation is discussed.

Motoneurons supplying the transverse mandibular muscle (TMM) in the guinea pig have been traced by injecting wheat germ agglutinin-horseradish peroxidase (WGA-HRP) in the TMM, and after applying HRP to the mylohyoid nerve. The TMM is bilaterally innervated by 22-36 motoneurons in each trigeminal motor nucleus, forming a column located ventrolaterally along the entire length of the superficial masseter motoneuron group. The axons are incorporated to the mylohyoid nerve. The location, the axon pathways in the brainstem and the pattern of the dentritic tree suggest that in the guinea pig the TMM motoneurons are involved in the masticatory movements in coordination with other jaw-closing muscles.

Injections of cholera toxin B subunit conjugated to horseradish peroxidase (CTB-HRP) were made into the lingual branch of the hypoglossal nerve in four species of finch in order to identify the innervation of the mechanoreceptors of the dermal papillae of the tongue, and simultaneously to determine the pattern of central projections of lingual hypoglossal afferents. The results showed that hypoglossal fibers innervate all the Herbst corpuscles and terminal cell receptors of the elaborately organized papillae of the dorsum of the tongue, of the shorter papillae in the ventral tongue, and the loose collection of Herbst corpuscles in the subpapillary region. Labelled fibers were also observed in the intralingual glands, in the intrinsic tongue muscles, and in the posterodorsal epithelium where they formed budlike structures. Retrogradely labelled cell bodies were located in the jugular ganglion and their central processes ascended and descended throughout the brainstem within the descending trigeminal tract (TTD). Terminal fields were observed within the dorsolateral part of the nucleus caudalis of TTD, predominantly ipsilaterally, and within the medial part of the dorsal horn of the first 4-6 cervical segments bilaterally. There were dense patches of termination over a dorsolateral subnucleus of the interpolated nucleus of TTD, and within two regions of the principal sensory trigeminal nucleus: a large one laterally and a small one medially. Terminal fields were also observed within the nucleus ventralis lateralis anterior of the rostral solitary complex, and within adjacent nuclei, which are probably equivalent to the dorsal sensory nuclei of the facial and glossopharyngeal nerves of other avian species. The results are interpreted in the light of the role of the tongue in species-specific patterns of feeding in finches, and the possible requirement for the central integration of touch and taste.

The projections of ventral medullary reticular neurons on both trigeminal (Vth) and hypoglossal (XIIth) motor nucleus were studied in sheep anesthetized with halothane. In a first series of experiments, extracellular microelectrodes were used to record the activity of medullary swallowing interneurons (SINs) located in the ventral region (around the nucleus ambiguus) of the swallowing center. Antidromic activation after electrical stimulation of the Vth and XIIth nuclei was tested in 83 SINs. For 38 SINs a clear antidromic activation was observed and for 8 of them the response was triggered by stimulation of either nucleus. As confirmed by the reciprocal collision test, these 8 SINs had branched axons sending information to both nuclei tested. Average latencies for antidromic activation of branched SINs after stimulation of the XIIth and the Vth motor nucleus were 2.2 +/- 0.6 ms and 2.7 +/- 0.8 ms respectively. The axonal conduction velocity of these neurons was 4.4 +/- 1.3 m/s for the collateral to the Vth motor nucleus and 2.7 +/- 0.7 m/s for axons projecting to the XIith motor nucleus. In a second series of experiments the double retrograde labeling technique was used to confirm the existence of neurons with branched axons in the medullary regions corresponding to the swallowing center. Small and well localized injections of Fast Blue (FB) and Diamidino Yellow (DY) fluorescent tracers were made in the Vth and in the XIIth motor nucleus respectively. A relatively large number of double-labeled cells was found in the ventral region of swallowing center (reticular formation around the nucleus ambiguus, 2-4 mm in front of obex).(ABSTRACT TRUNCATED AT 250 WORDS)

Recordings were made from convergent neurons in trigeminal nucleus caudalis of the rat. These neurons could be activated by both innocuous and noxious mechanical stimuli applied to their excitatory receptive fields on the ipsilateral part of the muzzle. Percutaneous application of suprathreshold, 2 ms square-wave electrical stimuli to the centre of the excitatory field resulted in responses to A- and C-fibres being observed. The effects on these responses of manual acupuncture performed by a traditional Chinese acupuncturist at the "Zusanli" point on the right hindlimb were compared with the effects induced by acupuncture applied at a non-acupoint, next to "Zusanli". In addition, the effects of acupuncture were compared with the inhibitory effects evoked by noxious thermal stimulation of the left hindlimb on the responses of the same neurons. This last type of inhibition has been described previously by our group and termed diffuse noxious inhibitory controls. Acupuncture, either applied at "Zusanli" or at a non-acupoint and noxious thermal stimulation induced similar strong inhibitory effects on the C-fibre-evoked responses of trigeminal convergent neurons (77.9 +/- 4.4%; 72.5 +/- 4.6% and 78.5 +/- 3.6% inhibition, respectively) and these inhibitions were followed by long-lasting aftereffects. In addition, both the acupuncture- and noxious thermal stimulation-evoked inhibitions were significantly reduced by systemic naloxone (0.4 mg/kg, i.v.). Since the antinociceptive effects elicited by acupuncture (i) had a similar magnitude and time-course to those evoked by noxious thermal stimulation, (ii) exhibited a lack of topographical specificity and (iii) involved an opioidergic link, we would suggest that, at least in our experimental conditions, acupuncture manoeuvres trigger the neuronal mechanisms involved in diffuse noxious inhibitory controls.

Horseradish peroxidase (HRP) was injected into either a single maxillary or a single mandibular primary (deciduous) cuspid tooth of 8- to 10-week-old kittens. The large apex of the primary cuspid allowed for some leakage of the HRP from the pulpal chamber to the periodontal ligament (PDL). Thus, the injection procedure resulted in the application of HRP to the PDL as well as to the pulpal tissues. The transganglionic transport of HRP resulted in discrete terminal fields within the spinal trigeminal nucleus (STN) and the main sensory nucleus (MSN). These projections were clearly somatotopically organized within the STN, but less so within MSN. Within pars oralis (PO) and pars interpolaris (PI), mandibular cuspid dental structures (MdCDS) were represented in a dorsal position relative to the maxillary cuspid dental structures (MxCDS), whereas within pars caudalis (PC) and the adjacent reticular formation the somatotopic representation was not dorsoventral, but rather mediolateral, with the MdCDS represented more medially than the MxCDS. Areas of overlap between MxCDS and MdCDS were found within MSN and to a lesser degree within the superficial laminae of PC. In addition, the fiber pathway leading to labeled somata in the mesencephalic trigeminal (Mes V) nucleus was clearly identified. The majority of the fibers traced to the Mes V nucleus exited the spinal trigeminal tract at the level of the transition from PO to the MSN and traversed the nuclear region in a position dorsal to and separate from the trigeminal motor tract. As in STN, fibers within the caudal Mes V tract appeared to be somatotopically organized, with the fibers from the MdCDS generally more dorsal than the ones from the MxCDS. Labeled fibers, some with terminal arbors, were also identified in close association with the trigeminal motor tract. The findings show a complex pattern of central representation in the immature feline central nervous system for deciduous dental structures.

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.

Retrograde transport and immunohistochemical double-labeling methods (Weinberg et al., 1985) were used to assess the distribution and projection status of spinal trigeminal (SpV) neurons that stain positively for glutamic acid decarboxylase (GAD) or gamma-aminobutyric acid (GABA). Large bilateral injections of diamidino yellow into the rostral and lateral pons, inclusive of V nucleus principalis and the parabrachial nucleus, retrogradely labeled large numbers of cells in each SpV subnucleus. Many cells in SpV subnuclei caudalis, interpolaris, and oralis also exhibited GABA immunoreactivity; the largest numbers were in caudalis and the smallest numbers were in oralis. However, none of the GABA- or GAD-immunoreactive SpV cells were double-labeled with diamidino yellow, though some reticular neurons displayed both GABA and the retrograde tracer. This negative result refutes a previously offered hypothesis that SpV local-circuit neurons with principalis collaterals are GABA-ergic (Jacquin et al., 1989b). These data also indicate that parabrachial-projecting SpV neurons are not GABA-ergic.

Motoneurons innervating the tensor tympani muscle were identified in the adult guinea pig and cat by the horseradish peroxidase (HRP) method. After HRP injection into the tensor tympani muscle, HRP-labeled neurons were seen in the regions outside the cytoarchitectonically-defined confines of the trigeminal motor nucleus; in the regions rostral to the rostral pole of the nucleus, as well as in the regions ventral and ventrolateral to the nucleus at the levels of the rostral half (guinea pig) or the rostral two-thirds (cat) of the nucleus. The tensor tympani motoneurons were generally smaller than the masticatory motoneurons.

In previous studies we have described vibrissae-related segmentation in the brainstem, thalamus, and cortex of the neonatal rat. Using succinic dehydrogenase (SDH) histochemistry, we delineated the time course in development of the normal segmentation at each level of the trigeminal system and the aberrant segmentation resulting from follicle damage at birth (Killackey and Belford, '79; Belford and Killackey, '79a, b). The present study examines the aberrant patterns that result from damage to the vibrissae follicles at different ages, comparing the patterns at the different levels of the trigeminal system. The present study indicates a number of similarities between the central representations of the vibrissae. First, the patterns are similar within each of the three representations in the trigeminal nuclei for removal at a given age in a particular animal. These changes include a decrease in SDH density but a maintenance of normal row widths. Second, the patterns are similar within both the ventrobasal complex and layer IV of somatosensory cortex for removal at a given age in a particular animal. These changes include a fusion of individual clusters into bands, and a decrease in band width, but maintenance of normal SDH density. Third, the effects of damage to a row of vibrissae follicles at different ages are graded. Earlier damage produces more marked aberrations. Fourth, for all of the structures, the transition between bands and clusters occurs with damage at the same age. Further, the last age at which damage produces aberrant patterns is Day 3 for all of the structures. Thus, the data suggest that there is one sensitive period for pattern alteration in the entire central trigeminal system.

In 22 adult sharks representing eight species, the mesencephalic trigeminal nucleus (Mes V) was examined employing the Romanes reduced silved method. Morphological evidence is presented of extensive development of neurites and intranuclear communication among Mes V cells of sharks, as well as their apparent innervation by extranuclear neurons. Species differences and the possible significance of the morphology of the Mes V in sharks to certain aspects of their behavior are discussed. A brief comparison of the Mes V of elasmobranchs and those of reptiles and mammals is included.

A combination of autoradiography and horseradish peroxidase histochemistry was used to identify telencephalic structures linking the sensory and motor components of the trigeminal system in the pigeon. A direct telencephalic projection from the principal trigeminal sensory nucleus upon the nucleus basalis via the quintofrontal tract was confirmed. Nucleus basalis projects upon a belt of neurons within the overlying neostriatum. This region (neostriatum frontale, pars trigeminale: NFT) gives rise to the fronto-archistriate tract which terminates both in the archistriatum intermedium and in the overlying neostriatum caudale, medial to the ventricle (neostriatum caudale, pars trigeminale: NCT). NCT projects, in turn, upon a region of archistriatum intermedium containing cell bodies of the occipito-mesencephalic tract. This pathway provides a link between the telencephalon and premotor areas within the lateral (parvicellular) reticular formation of the lower brainstem. The trigeminal sensorimotor circuit defined in these experiments has been implicated by neurobehavioral studies in the control of pecking, grasping, and feeding in the pigeon.

Following large injections of horseradish peroxidase into the thalamus of the adult rat, labeled neurons can be detected in the contralateral principal sensory nucleus and the subnucleus interpolaris of the spinal trigeminal nucleus. Thalamic projection neurons in the principal sensory nucleus are arranged in discrete aggregates which replicate the organization of the vibrissae and other parts of the face. This cellular organization corresponds very well with the pattern of vibrissae related afferent termination in this region of the nucleus as seen in SDH preparations. There is no such cellular organization in the subnucleus interpolaris.

The retrograde transport of horseradish peroxidase (HRP) was used to assess the overall morphology of neurons in the dorsal half of the border zone (BZ) of rat trigeminal nucleus oralis (Vo) that project to ipsilateral orofacial portions of four major tactile areas (crura I and II, the paramedian lobule, and the uvula) of the cerebellar cortex. We wished to answer two important questions: Does this group of cells consist of one or more morphologically distinct types, each projecting to different cerebellar tactile areas? Does the overall morphology or morphologies of these BZ neurons resemble one or more of the five types of identified trigeminocerebellar neurons in the dorsomedial (DM) subdivision of Vo, or do these trigeminocerebellar cells represent an additional morphologically distinct type or types restricted to BZ? The morphology of BZ neurons innervating the orofacial portions of all four cerebellar tactile areas was similar and did not resemble that of any of the five types of DM cells. They were characterized by a pyramidal- to fusiform-shaped cell body measuring 10-13 X 20-25 microns, which emitted three or four primary dendrites; one was directed dorsally, while the others took a more ventral trajectory. The primary dendrites generated a dendritic arbor arranged as a flattened disk oriented parallel to the spinal V tract. The dendritic field was largely restricted to BZ; it measured up to 150 microns in width, and spanned up to 450 microns dorsoventrally and rostrocaudally. An axon arose from the dorsal aspect of the cell body and gave rise to a single short collateral within 10 microns of its origin. This collateral remained unbranched and generated several boutons within BZ, while the parent axon, without branching further, traveled dorsolaterally toward the inferior cerebellar peduncle. Frequently, a second axon arose ventrally from the soma, and after a short unbranched course entered a deep axon bundle, where it assumed a descending trajectory. The intranuclear portion of this second axon was characterized by several boutons en passant.

The mesencephalic trigeminal nucleus was studied in anaesthetized and curarized rabbits by recording the unitary activity through extracellular microelectrodes and identifying the constituent cell types. Two types of units were found, namely primary afferents supplying jaw raising muscle spindles and periodontal or gingival mechanoreceptors. These two groups of neurons exhibited a rostrocaudal somatotopy: the former occupied the entire rostral portion of the nucleus (A7-P2.3; trochlear decussation being taken as an arbitrary 0 level), the latter was located caudally (P3-P4.5) while the somata of both types of afferent fibres were present between P2.2 and P3. No evidence was found for representation of both tendon organs of jaw muscles and joint receptors. Among the units innervating muscle spindles, secondary afferents were largely more numerous than the primary ones. Among periodontal and gingival mechanoreceptor afferents, incisors were the most widely represented, followed by interalveolar gingiva and molars; the axonal conduction velocity ranged between 9 and 40 m/sec and between 8 and 16 m/sec for ipsilaterally and contralaterally projecting neurons, respectively. The motor responses obtained by electrical stimulation of discrete areas of the MTN confirmed the presence of a high degree of segregation between the two different populations of neurons. In fact, jaw raising movements are obtained when stimulating the area within A7 and P2 containing the somata of spindle afferent neurons, while only jaw opening movements are elicited by stimulation of the caudal levels of the nucleus. These data also show that the periodontal neurons whose somata are located in the MTN participate in the jaw opening reflex, just as the more numerous periodontal mechanoreceptors whose somata are located in the Gasser ganglion. Soma-somatic and soma-axon hillock gap junctions were found among the neurons of the MTN, particularly in the caudal third of the nucleus.

The efferent and intranuclear connections of the nuclei of the descending trigeminal tract of the mallard have been studied with lesion methods, and by axonal transport techniques following injections of tritiated leucine, and of horseradish peroxidase. The large subnucleus oralis neurons, including those belonging to the nucleus of the ascending glossopharyngeal tract, have proven to be the sole origin of trigeminocerebellar connections. The cerebellar afferents are of the mossy fiber type, and terminate predominantly in lobules V, VI and VII, and possibly, lobule IV. Trigeminocerebellar projections are ipsilateral except for the vermal area. Subnucleus interpolaris is the main source of intratrigeminal fibers that terminate in subnucleus oralis and the ventral part of the main sensory nucleus. These intranuclear connections are bilateral, but the medium-celled caudal part of subnucleus interpolaris in particular contains the majority of bi- and/or contralaterally projecting neurons. Additionally, the small cells in the rostral part of subnucleus interpolaris project ipsilaterally upon the parabrachial region, and upon the lateral reticular formation. Projections upon the parabrachial region furthermore emanate bilaterally from layer I of the rostral subnucleus caudalis. A minor part of layer I neurons sends its axons contralaterally along with those of the dorsal column nuclei toward the thalamic nucleus dorsolateralis posterior. Associated with the medial lemniscus, contralateral termination is also present in the lateral part of the ventral lamella of oliva caudalis, in the marginal zone of nucleus mesencephalicus lateralis, pars dorsalis and immediately surrounding intercollicular grey and, finally, in the nucleus intercalatus thalami. Furthermore, a bilaterally descending projection from subnucleus caudalis upon layers I and II of the rostral cervical cord was observed. Close to their origin subnucleus caudalis neurons project upon the adjoining caudal part of the lateral reticular formation.

Using the retrograde horseradish peroxidase technique, we have examined the distribution of labeled thalamic-, spinal- and cerebellar-projecting neurons in the trigeminal sensory nuclei of the cat. Injections into the nucleus ventralis posterior of the thalamus resulted in labeling of neurons in lamina I (subnucleus zonalis), the deeper part of lamina IV (the subnucleus magnocellularis) of the nucleus caudalis and in lamina V (the lateral extension of the nucleus medullae oblongatae centralis) on the contralateral side. A very large number of labeled small neurons were observed mainly in the caudal part of the nucleus interpolaris and in the ventral division of the principal sensory nucleus on the contralateral side and in the dorsal division of the principal sensory nucleus on the ipsilateral side. Injections into the known projection areas of the cerebellar cortex labeled mainly ipsilaterally the trigeminocerebellar neurons in a restricted ventrolateral area of lamina IV of the nucleus caudalis at its rostral level and in lamina V. Many labeled neurons were also observed in the nucleus interpolaris. Although the distribution overlapped with that of the trigeminothalamic neurons, the greatest majority were concentrated in its rostral part where the trigeminothalamic neurons were very small in number. In addition, labeled neurons were observed in the rostral part of the nucleus oralis and the ventralmost part of the ventral division of the principal sensory nucleus. No labeled neurons were observed in the dorsal division of the principal sensory nucleus and the mesencephalic nucleus. The trigeminospinal neurons were labeled mainly ipsilaterally following injections into the upper cervical cord. They were located in laminae I and III, the deeper part of lamina IV of the nucleus caudalis and in lamina V. Only scattered labeled neurons were found in the nucleus interpolaris. The number of labeled neurons increased in the nucleus oralis at the level of the superior olive. They tended to be distributed around or dorsal to the group of the trigeminothalamic neurons at the caudal part of the principal sensory nucleus. No neurons of the principal sensory nucleus appeared to project to the spinal cord. Based on the large size and location, the trigeminospinal neurons could be differentiated from the other projection neurons in the nucleus oralis. The present study demonstrates that the trigeminal sensory nuclei are composed of groups of neurons with different projections, since the main aggregations are localized at different levels. However, it should be examined whether the neuronal groups, which are labeled from the different structures in similar locations, are composed of individual neurons projecting to more than one of these structures.

The central projections of afferent fibers from the cornea, and the infraorbital, infratrochlear, frontal, lacrimal and auriculotemporal nerves were investigated by means of the transganglionic transport of horseradish peroxidase. Afferent projections to the dorsal horn of the medulla are organized along both the rostrocaudal axis and the ventrolateral to dorsomedial margin of the medullary dorsal horn. An inverted but discontinuous facial representation exists through the restrocaudal axis of the dorsal horn of the medulla with perioral and nasal receptive fields innervated by the infratrochlear nerves represented rostral to the progressively more posterior receptive fields innervated by the frontal, lacrimal and auriculotemporal nerves, respectively. The organization of the primary afferents is not uniform over the laminae of the dorsal horn of the medulla; the projections from the different nerves show the least overlap in lamina II, while overlap is most extensive in laminae I and V. The sensory projection from the cornea to the medullary dorsal horn is most dense in laminae I and II. All nerves, including those innervating the cornea, project to the interpolar, oral and principal trigeminal nuclei and are somatotopically organized. Projections to the reticular formation and the contralateral trigeminal sensory complex were not found in this study. These results support the organization of the dorsal horn of the medulla proposed by Déjerine ('14) and show that this organization is most evident for the primary afferent projections to lamina II.

Horseradish peroxidase (HRP), injected intramuscularly, specifically labeled motoneurons innervating antagonistic jaw muscles in the cottonmouth mocassin, Agkistrodon piscivorus piscivorus. Adductor mandibulae profundus, part 3a, motoneurons were localized in the lateral regions of the ventral and intermediate subnuclei of the trigeminal (V) motor nucleus. These were large cells containing fine, granular reaction product characteristic of alpha-motoneurons. Small cells, which contained large coarse reaction granules characteristic of gamma-motoneurons were localized in a separate cluster in the lateral regions of the dorsal subnucleus of the V motor nucleus. Depressor mandibulae motoneurons were localized in the ventromedial regions of the facial (VII) motor nucleus, primarily in the caudal half. Cell sizes ranged from 30--50 micrometers in diameter and HRP staining characteristics were variable, indicating a mixed population of motoneuron functional types without the segregation noted in the V motor nucleus. Boutons which made synaptic contact with labeled somata or processes were classified according to morphological type and their frequency of appearance. Boutons containing spherical vesicles (S-, C-, T-) were distributed similarly on motoneurons of both muscles, but more F-boutons, those with flattened vesicles, synapsed on the adductor motoneurons. Comparison of snake bouton distribution with that in mammalian spinal cord indicates that synaptology on all these motoneurons are remarkably similar. The more frequent occurrence of C-boutons, those with a subsynaptic cistern, on reptilian motoneurons may indicate a stronger intrasegmental input, as determined from mammalian degeneration studies.

Young adult albino rats of Wistar strain were used for the present study. 0.5 to 15 microliters of 20-50% of horseradish peroxidase (HRP) were injected into each individual muscle of mastication to label neurons in the trigeminal motor nucleus (TMON) for light microscopic study. The results reveal that: (1) Many HRP-labeled, multipolar neurons are observed in the motor nucleus in each jaw-closing muscle (JCM) with less in each the jaw-opening muscle (JOM). (2) The motor neurons innervating each masticatory muscle in the motor nucleus show a somatotopic arrangement: (a) those innervating the temporalis muscle are located in the medial and dorsomedial parts; (b) those innervating the masseter muscle are located in the intermediate and lateral; (c) those innervating the medial and lateral pterygoid muscles are located in the lateral, ventrolateral and ventromedial parts, respectively; and (d) those innervating the mylohyoid and the anterior belly of the digastric muscles are located in the most ventromedial part of the caudal one-third of the nucleus. Axons of most masticatory motor neurons run ventrolaterally in between the motor and the chief sensory nuclei of the trigeminal nerve. However, those of the mylohyoid and anterior belly of the digastric muscles ascend dorsally to the dorsal aspect of the caudal nucleus and then turn ventrolaterally to join the motor root of the trigeminal nerve. Furthermore, the dendrites of the motor neuron of JCM converge dorsocaudally to the supratrigeminal region. The diameters of neurons of each JCM display a bimodal distribution. However, an unimodal distribution is present in the motor neurons from each JCM. It is suggested that the motor nucleus innervating the JCM is comprised of comprised of alpha- and gamma-motor neurons. It, thus, may provide a neural basis for the regulation of the muscle tone and biting force.

This study demonstrates that trigeminal nucleus oralis, the most rostral subdivision of the spinal trigeminal nucleus, contains four morphologically distinct types of small neurons which project to the medullary dorsal horn (trigeminal nucleus caudalis) via descending intratrigeminal pathways. Using the retrograde transport of horseradish peroxidase following injections in the medullary dorsal horn, labeled small neurons with cell bodies ranging from 8-15 microns in diameter are found principally in the ventrolateral portion of the trigeminal nucleus oralis. Most neurons are labeled ipsilaterally throughout the entire rostrocaudal extent of the ventrolateral portion of the trigeminal nucleus oralis, but a few cells are also labeled contralaterally. From this aspect of the present study it can be concluded that a specific portion of the trigeminal nucleus oralis, i.e. the ventrolateral part, contains numerous small neurons which send descending projections to the medullary dorsal horn that could affect synaptic activity there. Utilizing both the methods of Golgi and retrograde horseradish peroxidase labeling four distinct types of small descending medullary dorsal horn projection neurons can be distinguished in the ventrolateral portion of the trigeminal nucleus oralis on the basis of their morphology and the distribution of their axons and dendrites. All four neuronal cell types are present throughout the entire rostrocaudal extent of the trigeminal nucleus oralis. Type I neurons are the most frequently labeled descending medullary dorsal horn projection neurons. They are concentrated in the medial 500-550 microns of the ventrolateral portion of the trigeminal nucleus oralis and display dendritic trees which occupy spherical domains approaching 300 microns in diameter. The unmyelinated axons of many of these cells arise either directly from the cell body or a primary dendrite and give rise to a single collateral within 50 microns of their site of origin. This collateral generates a fine axonal plexus within a portion of the dendritic arbor of the parent cell while the parent axon, without branching further, travels a short distance in the ventrolateral portion of the trigeminal nucleus oralis and enters a deep axon bundle. Type II neurons are the second most frequently labeled descending medullary dorsal horn projection neuron. They generate medial and lateral dendritic arbors which together span nearly the entire medial 500-550 microns of the ventrolateral portion of the trigeminal nucleus oralis. An unmyelinated axon emerges from the cell body and within 10-30 microns of its origin gives rise to two collaterals.(ABSTRACT TRUNCATED AT 400 WORDS)

The spinal projections from the mesencephalic trigeminal nucleus (Probst's tract) were investigated in the duck by HRP method. From our researches it results that the caudal extension of the Probst's tract reaches the T1 segment of the spinal cord.

The retrograde horseradish peroxidase technique was used to locate neurons projecting to rostral trigeminal areas via ascending intranuclear pathways. After rostral trigeminal injections, labeled neurons were found in the subnucleus interpolaris and in all laminae of the medullary dorsal horn. Most neurons were labeled ipsilaterally in laminae III and IV, but some small neurons were labeled in lamina II in the medullary dorsal horn. The distribution of labeled cells suggested that these projections are topographically arranged. Labeled neurons were found in lamina I and lamina V, bilaterally; this was especially true if the parabrachial complex was included in the injection.

The anatomical connections between two regions involved in swallowing, the medullary reticular formation around the nucleus ambiguous (ventral group of swallowing interneurons) and the trigeminal motor nucleus (NMV) were studied in sheep using anterograde and retrograde tracing techniques. [3H]leucine and HRP were injected into the ventral medulla and the NMV respectively. These injections were performed according to stereotaxic coordinates previously established by electrophysiological recording tests. The results show bilateral connections between the ventral medulla, 2-4 mm rostral to the obex, and the trigeminal motor nucleus. In addition, the same area of the ventral medulla is connected with the homologous contralateral region and with the facial, vagal and hypoglossal nuclei that are also involved in swallowing. Some connections between the ventral medulla and the NMV must be monosynaptic since aggregates of silver grains typical of axon terminals were observed on the cell bodies of trigeminal motoneurons. Obviously, the various connections shown to exist in this study may serve other functions than swallowing. Nevertheless, most of the results are consistent with the view that the ventral group of medullary swallowing interneurons are switching neurons that distribute the programmed swallowing excitation to the various motoneuron pools involved in deglutition.

Horseradish peroxidase was injected under pressure in the main excretory duct of the submandibular gland of cats, to locate the perikarya of the preganglionic parasympathetic salivatory neurons in the brainstem. Labeled multipolar neurons were found ipsilaterally in the lateral reticular formation, where they extended from the rostral plane that contains the genu of the facial nerve to the caudal plane that contains the rostral pole of the dorsal motor nucleus of the vagus nerve.

Following horseradish peroxidase injection into the cervical and lumbosacral cords of adult albino rats, labeled neurons were seen in the first cervical segment, brain stem, and cerebellar and diencephalic nuclei. A new pathway, the faciospinal projection, originating in the medial portion of the rostral part of the facial nucleus, was traced. Another new pathway, the olivospinal pathway, is probably also present. Our results for neurons projecting to the spinal cord (spinal-projecting neurons) from the nucleus ambiuus, dorsal motor nucleus of the vagus, superior vestibular nucleus and nucleus f, nucleus Darshevch, nucleus Rolleri, nucleus prepositus hypoglossi, and nucleus of the posterior commissure have been reported before in other mammals but not in rats. Projections from the following regions are in general agreement with previous results in rats, but show significant topographical differences: the first cervical segment; nuclei gracilis, cuneatus, and cuneatus lateralis; the midline and lateral reticular nuclear complex; the trigeminal nuclear complex (spinal, principal, and mesencephalic); nucleus of the tractus solitarius; the medial, lateral, and descending vestibular nuclei, nuclei coeruleus and subcoeruleus; superior colliculus; interstitial nucleus of Cajal, and the deep cerebellar nuclei. The distribution of labeled neurons in the nucleus parabrachialis, nucleus tegmentolaterodorsalis, nucleus Kölliker-Fuse, nucleus Edinger-Westphal, and the hypothalamic nuclear complex confirmed that of previous reports in rats. With the exception of a few nuclear groups which project primarily to either lumbosacral (e.g., the paraventricular nucleus of the hypothalamus) or cervical segments (e.g., the facial motor nucleus and the superior colliculus) most of the other nuclear groups project to both the lumbar and cervical levels. There is no distinct somatotopy in the neuronal groups projecting to both cervical and lumbosacral levels. With only a few exceptions (e.g., the superior vestibular nucleus) most of the spinal-projecting neurons are bilaterally distributed, some with contralateral and others with ipsilateral predominance.

Horseradish peroxidase was injected into various levels of the spinal cord of turtles (Pseudemys and Chrysemys), lizards (Tupinambis, Iquana, Gekko, Sauromelus, and Gerrhonotus), and a crocodilian (Caiman). The results suggest that brainstem reticulospinal projections in limbed reptiles rival mammalian reticulospinal systems in complexity. The reptilian myelencephalic reticular formation can be divided into four distinct reticulospinal nuclei. Reticularis inferior pars dorsalis (RID) contains multipolar neurons which project bilaterally to the spinal cord. Reticularis inferior pars ventralis (RIV), which is only found in lizards and crocodilians, contains fusiform neurons with horizontally running dendrites and it projects ipsilaterally to the spinal cord. Reticularis ventrolateralis (RVL), which is found only in field lizards, contains triangular neurons whose dendrites parallel the ventrolateral edge of the brainstem and it projects ipsilaterally to the spinal cord. The myelencephalic raphe (RaI) varies considerably. RaI of turtles contains large reticulospinal neurons which form a continuous population with more laterally situated RID cells. RaI of lizards contains a few small reticulospinal neurons. RaI of the crocodilian Caiman contains giant reticulospinal neurons with laterally directed dendrites. The caudal metencephalic reticular formation of reptiles can be divided into two distinct reticulospinal nuclei. Reticularis medius (RM) contains large neurons with long, ventrally directed dendrites; it projects ipsilaterally to the spinal cord. Reticularis medius pars lateralis (RML) contains small neurons with laterally directed dendrites; it projects contralaterally to the spinal cord. The rostral mesencephalic and caudal mesencephalic reticular formation of reptiles can be divided into three distinct reticulospinal nuclei. Reticularis superior pars medialis (RSM) consists mostly of small, spindle-shaped neurons which project bilaterally to the spinal cord. In the lizard Tupinambis, however, large multipolar, ipsilaterally projecting neurons are occasionally seen in RSM. Reticularis superior pars lateralis (RSL) contains large, ipsilaterally projecting neurons with long, ventrolaterally directed dendrites. SRL in lizards can be divided into a dorsomedial portion, which projects ipsilaterally to the spinal cord, and a ventrolateral portion which projects contralaterally. The locus ceruleus-subceruleus field (LC-SC) contains small spindle-shaped neurons which project bilaterally to the spinal cord. Labelled reticulospinal neurons were also observed in the rostral metencephalic raphe (RaS) of the turtle brainstem. These cells are small, spindle-shaped neurons which resemble the small cells of the adjacent RSM field.

While the somatotopic organization of many central systems is well characterized, that of peripheral sensory neurons has not been adequately defined. This is especially true for the trigeminal ganglion. By applying HRP subcutaneously at each of 14 sites and also intramuscularly, it is possible to determine whether the location of sensory neurons within the ganglion reflects their peripheral projections. There is no discernible somatotopic organization of neurons in the ophthalmic lobe. However, the location of maxillary neurons in the maxillo-mandibular lobe is organized with the most posterior cells projecting to sites closest to the ganglion and with neurons located more anteriorly projecting to progressively more distant sites. There is a less well defined organization in the superior-inferior axis of the ganglion, and none along its proximal (root) to distal axis. Mandibular exteroceptive neurons are found primarily in the anterior region of the maxillo-mandibular lobe, while mandibular proprioceptive cells are located in the proximo-central part of this lobe. In most cases there is a considerable scattering of horseradish peroxidase (HRP)-filled neurons. Projections to contralateral ganglia, the trigeminal motor nucleus, and the trigeminal mesencephalic nucleus were also examined. Cytologically, the hatchling trigeminal consists of two interspersed types of neurons: large, lightly staining and smaller, darkly staining cells. Previous experiments have proved that these two cell types do not correspond to each of the embryonic precursors of trigeminal neurons, the neural crest and placodal cells. In this study HRP was found localized in both classes of neurons following injection at all sites, including jaw-closing muscles. This indicates that the dual cytology is not correlated with either distribution of peripheral fibers or exteroceptive vs. proprioceptive functions. The possibilities that the two types of neurons may have different central projections and/or may be related to visceral vs. somatic afferent functions are discussed.

Central pathways converging upon the trigeminofacial motor nuclei of the mallard were studied in order to elucidate neuroanatomically the presumed influence of primary sensory trigeminal afferents upon jaw muscle activity. The techniques used included the Fink-Heimer I method after lesions, and axonal transport labeling following injections of 3H-leucine or of HRP for retrograde identification of the neurons of origin. A general description is given of the trigeminofacial motor complex. Jaw closer muscles are innervated by trigeminal motor neurons, and facial motor neurons innervate the jaw depressor muscles. Two afferents premotor systems, one including the mesencephalic trigeminal nucleus (MesV) and the other the rhombencephalic reticular formation, are distinguished. The proprioceptive neurons of the mesencephalic trigeminal nucleus project upon the ipsilateral trigeminal motor nucleus and upon the nucleus supratrigeminalis. The latter cell group bilaterally projects upon the dorsal and intermediate parts of the facial motor nucleus and upon the dorsal and intermediate parts of the facial motor nucleus and upon part of the trigeminal motor nucleus. Exteroceptive information, relayed through the primary sensory trigeminal column (PrV and nTTD), ultimately reaches the motor nuclei via the reticular formation. The reticular formation forms the final link of three separate circuits: a telencephalic one entered through the principal trigeminal sensory nucleus, a cerebellar one via subnucleus oralis of the descending trigeminal system, and a direct one via subnucleus interpolaris. No direct connections between the principal trigeminal sensory nucleus or subnuclei of the descending trigeminal system and the motor nuclei of the trigeminal (NV) and facial (NVII) nerves have been observed, nor are such direct projections present in the outflow of the presumed telencephalic and cerebellar circuits, viz. of the archistriatum and the central cerebellar nuclei, respectively. The archistriatum projects via the occipitomesencephalic tract upon the lateral rhombencephalic reticular formation as far down as the rostral cervical cord, as well as upon the subnucleus interpolaris of the descending trigeminal system. Similarly, efferents from the central cerebellar nuclei reach the reticular formation, which in turn projects bilaterally upon the motor nuclei. Finally, commissural intermotor connections apparently are mediated by reticular cells surrounding the motor nuclei of NV or NVII, rather than emanating from these nuclei directly.

Some headaches of organic nature can be readily attributed to excitation of nerve terminals or receptors located in distinct intracranial structures. The trigeminal system mediates most of the intracranial pain sensation. Modern tracer methods have investigated the somatotopic organization of meningeal sensation in the Gasserian ganglion. The dura mater is supplied by the trigeminal divisions sectorially, in a similar manner to the face. Although the notion of somatotopic representation of the meninges in the Gasserian ganglion may suggest possible surgical intervention for intractable headache at the level of the Gasserian ganglion, separation of the facial and corneal representation from the meningeal cell pool is not sharp enough to avoid facial dysesthesias.

Transganglionic transport of horseradish peroxidase (HRP) has been used to study the cell bodies and central projections of neurons innervating the vibrissae in the rat. These can be grouped into five horizontal rows and one posterior vertical row. Twenty-four to 48 hours after the nerves innervating different vibrissae were exposed to HRP, the trigeminal ganglia, brainstem, and upper cervical spinal cord were fixed by perfusion and serial sections were processed according to the tetramethylbenzidine technique. The results revealed a tendency for somatotopic organization in the trigeminal ganglion of cell bodies innervating the different vibrissae. Corresponding termination areas in the trigeminal sensory nuclei showed a detailed pattern of organization replicating the peripheral organization of the vibrissae. In all trigeminal sensory nuclei the horizontal rows are represented in an inverted fashion from dorsal to ventral, i.e., the most dorsal row is represented most ventrally. In addition, the more anterior a vibrissa is located, the deeper is it represented in the rostral nonlaminated nuclei. The situation is reversed in the laminated nucleus caudalis. The posterior vertical row is represented most superficially in the rostral nonlaminated nuclei, but most deeply in the laminated nucleus caudalis. In nucleus caudalis there are also rostrocaudal differences in the representation of different vibrissae. Thus, the posterior vibrissae in a horizontal row have their main representations more caudally than the anterior vibrissae. The posterior vertical row has its main representation most caudally, in the C1 segment.

The cobalt labeling technique was applied to the trigeminal, facial, and abducens roots in order to investigate the location and morphology of motoneurons of the main and accessory motor nuclei of these cranial nerves in the rat. On the basis of soma-dendritic morphology, axonal trajectory, and tegmental location, three different accessory nuclei were distinguished in the brainstem. Neurons of the accessory nuclei of V and VII form a common club-shaped nucleus extending between the main motor nuclei of V and VII. The dendritic tree of these neurons is oriented in the dorsoventral direction and the axon invariably follows a dorsolaterally directed bent course. The accessory VI nucleus is located ventral and lateral to the main VI nucleus in the lateral tegmentum. The neurons have elongated perikarya, strong ventral and weak dorsal dendritic arbors, and the bent axonal course is directed dorsomedially. Neurons of the main VI nucleus have rounded perikarya and radial dendritic arborizations. Muscle groups innervated by the accessory and main motor nuclei have distinctly different functions.

The horseradish peroxidase (HRP) retrograde transport method was used to identify brainstem afferents to the cerebellar flocculus in the pigmented rat. Injections of the enzyme were made through recording microelectrodes, making it possible to localize the injection site by physiological criteria. Clearly, the largest number of afferents arise from the bilateral vestibular and perihypoglossal nuclei and from the contralateral dorsal cap (of Kooy) of the inferior olive. Additionally, a substantial number arise bilaterally from: (1) the nucleus reticularis tegmenti pontis (NRTP); (2) several of the cranial motor nuclei including the abducens, retrofacial and facial nuclei and the nucleus ambiguus; (3) the rostral part of the lateral reticular nucleus (subtrigeminal nucleus); (4) the raphe pontis and raphe magnus and (5) neurons intercalated among the medial longitudinal fasciculus (MLF) just rostral to the hypoglossal nucleus and another group rostral to the abducens nucleus. The basilar pontine nuclei contained a large number of lightly labeled neurons in all flocculus injections which were discretely located within the dorsolateral, lateral and medial divisions. These areas were labeled bilaterally but with a slight contralateral preponderance. Injection into the flocculus, but involving the adjacent ventral paraflocculus, produced a heavier labeling of pontine neurons with a slightly different distribution. Therefore, we tentatively conclude that the flocculus receives input from these pontine visual centers (dorsolateral, lateral and medial nuclei), perhaps through collateral projections from neurons projecting to the paraflocculus. The present study demonstrates strong similarities between the rat and other species studied (e.g., rabbit, cat, monkey) in terms of the brainstem nuclei projecting to the flocculus. Most noticeable in quantitative terms are the pathways known to mediate vestibular (vestibular and perihypoglossal nuclei) and visual (optokinetic) information (e.g., NRTP). Additionally, we can provide morphological evidence that the midline and paramedian pontine tegmentum, identified in the cat and monkey as containing saccade-related neurons, send large numbers of projections to the rat flocculus. Given these similarities, the rat may be a suitable animal model in which to study the pathways underlying visual-vestibular interaction and saccadic mechanisms in the flocculus.

Descending spinal projections were investigated in the turtle Pseudemys scripta elegans following injections of horseradish peroxidase and/or radioactive wheat germ agglutinin into the spinal cord at various levels. Using various planes of section the cells of origin in the brainstem, cerebellum, and diencephalon were characterized according to their size, dendritic tree, and precise location. Projections to levels as far caudal as the lumbar spinal cord were found to arise from medial and lateral rhombencephalic reticular fields, including the perihypoglossal complex, the nucleus raphe inferior, and the locus coeruleus; from certain subdivisions of the vestibular complex (ipsilateral subnucleus (subn.) ventrolateralis, contralateral subn. ventromedialis, and possibly subn. tangentialis); from the motor trigeminal nucleus; from the contralateral red nucleus, the ipsilateral nucleus (n.) interstitialis of the fasciculus longitudinalis medialis (flm) and from the hypothalamus. Fibers to high cervical levels arose from neurons within the dorsolateral and superior vestibular nuclei, the lateral cerebellar nucleus, the mesencephalic trigeminal nucleus, and from neurons of the optic tectum. Low cervical and thoracic spinal levels were reached by fibers from the torus semicircularis n. laminaris, the n. of the flm, the medial cerebellar nucleus as well as from the n. vestibularis inferior and the principal and descending trigeminal nuclei.

Toxic ricin was used to study the central distribution of dental afferents in the cat. Following intrapulpal ricin injections ganglion cell degeneration is seen in the II and III ganglion divisions. Central argyrophilic degeneration occurs in the dorsal portion of all ipsilateral trigeminal nuclei. Ventral degeneration is seen in the pars interpolaris and pars caudalis. No contralateral degeneration was observed. The results are discussed with regard to previous studies of the central location of dental afferents.