Cortical projections to nuclei adjacent to the oculomotor complex in the medial dien-mesencephalic tegmentum in the monkey

Cortical projections to cell groups surrounding the oculomotor complex were studied by using the retrograde and anterograde capabilities of the horseradish peroxidase (HRP) technique in old and new world monkeys. Fluid HRP injections or transcannular solid polyacrylamide HRP gel implants were made into the oculomotor nucleus (OMN) and adjacent nuclei to label retrogradely corticofugal neurons that project to this region, and cortical HRP gel implants were made in various areas of the frontal lobe to label anterogradely the trajectories and terminations of cortico-paraoculomotor projections and thus to confirm the retrograde findings. Projections to the paraoculomotor cell groups in the medial dien-mesencephalic tegmentum originate almost exclusively from the frontal lobe. Both retrograde and anterograde studies confirmed that the prearcuate cortex in the concavity of the arcuate sulcus, including the frontal eye field, and, to a lesser extent, suprarcuate rostral dorsal area 6 cortex and the dorsomedial convexity (area 9), project to the rostral interstitial nucleus of the medial longitudinal fasciculus (riMLF) in the dorsal region of the prerubral field, nucleus of Darkschewitsch (ND), medial accessory nucleus of Bechterew (NB) and dorsomedial parvocellular red nucleus (dmPRN). The premotor area 6 and motor area 4 cortex, on the other hand, give rise to projections that target a larger portion of the parvocellular red nucleus, extending rostrally into the ventral region of the prerubral field, and a rather intense projection to the ND. The interstitial nucleus of Cajal (IC) was distinguished more by its light, or lack of, projections from the frontal cortex. The inferior parietal lobule (IPL, area 7) which has certain common physiological properties with the frontal eye field (FEF area 8) related to the oculomotor system, lacked retrogradely labeled neurons in all cases where transcannular gel implants into the OMN eliminated the possibility of HRP uptake in the corpus callosum or other structures traversed in needle injections, suggesting that the IPL affects eye movement primarily through its rostrally directed corticocortical associational connections with the FEF. In additional cases, the ND-NB-dmPRN configuration of cells that receives FEF input is shown to project to the inferior olivary complex (i.e., is pre-olivo-cerebellar), whereas riMLF and IC give rise to descending projections in the MLF, which target extraocular muscle motor nuclei, vestibular complex, and spinal cord. The results are discussed in terms of the potential role of the cerebral cortex in eye movement mechanisms.     

Organization of cerebellar and area "y" projections to the nucleus reticularis tegmenti pontis in macaque monkeys

Axonal projections to the nucleus reticularis tegmenti pontis (RTP) were studied in 11 macaque monkeys by mapping axonal degeneration from lesions centered in the dentate and interpositus anterior (IA) nuclei and by mapping anterograde transport of tritiated amino acid precursors injected into the dentate nucleus. Projections from the dentate and IA nuclei overlap in central parts of the body of RTP, but the terminal field of dentate axons extends dorsomedial and rostral to the terminal field of IA axons, and IA terminal fields extend more ventrolaterally. A caudal to rostral topography of projections from each nucleus onto dorsal to ventral parts of RTP was seen. Projections from rostral parts of both nuclei terminate in a sublemniscal part of the nucleus. The topography of dentate and IA projections onto central to ventrolateral RTP appears to match somatotopic maps of these cerebellar nuclei with the somatotopic map of projections to RTP from primary motor cortex. Projections from caudal and ventral parts of the dentate nucleus appear to overlap oculomotor inputs to rostral, dorsal, and medial RTP from the frontal and supplementary eye fields, the superior colliculus, and the oculomotor region of the caudal fastigial nucleus. Projections to the paramedian part of RTP from vestibular area "y" were also found in two cases that correlated with projections to vertical oculomotor motoneurons. The maps of dentate and IA projections onto RTP correlate predictably with maps of dentate and IA projections to the ventrolateral thalamus and subnuclei of the red nucleus that were made from these same cases (Stanton [1980b] J. Comp. Neurol. 192:377-385).     

Periaqueductal gray matter projections to midline and intralaminar thalamic nuclei of the rat

The periaqueductal gray matter (PAG) projections to the intralaminar and midline thalamic nuclei were examined in rats. Phaseolus vulgaris-leucoagglutinin (PHA-L) was injected in discrete regions of the PAG, and axonal labeling was examined in the thalamus. PHA-L was also placed into the dorsal raphe nuclei or nucleus of Darkschewitsch and interstitial nucleus of Cajal as controls. In a separate group of rats, the retrograde tracer cholera toxin beta-subunit (CTb) was injected into one of the intralaminar thalamic nuclei-lateral parafascicular, medial parafascicular, central lateral (CL), paracentral (PC), or central medial nucleus-or one of the midline thalamic nuclei-paraventricular (PVT), intermediodorsal (IMD), mediodorsal, paratenial, rhomboid (Rh), reuniens (Re), or caudal ventral medial (VMc) nucleus. The distribution of CTb labeled neurons in the PAG was then mapped. All PAG regions (the four columns of the caudal two-thirds of the PAG plus rostral PAG) and the precommissural nucleus projected to the rostral PVT, IMD, and CL. The ventrolateral, lateral, and rostral PAG provided additional inputs to most of the other intralaminar and midline thalamic nuclei. PAG inputs to the VMc originated from the rostral and ventrolateral PAG areas. In addition, the lateral and rostral PAG projected to the zona incerta. No evidence was found for a PAG input to the ventroposterior lateral parvicellular, ventroposterior medial parvicellular, caudal PC, oval paracentral, and reticular thalamic nuclei. PAG --> thalamic circuits may modulate autonomic-, nociceptive-, and behavior-related forebrain circuits associated with defense and emotional responses.     

Organization of projections from the juxtacapsular nucleus of the BST: a PHAL study in the rat

The axonal projections of the juxtacapsular nucleus of the anterior division of the bed nuclei of the stria terminalis (BSTju) were examined with the Phaseolus vulgaris-leucoagglutinin (PHAL) method in the adult male rat. Our results indicate that the BSTju displays a relatively simple projection pattern. First, it densely innervates the medial central amygdalar nucleus and the subcommissural zone and caudal anterolateral area of the BST — cell groups involved in visceromotor responses. Second, it provides inputs to the ventromedial caudoputamen (CP) and anterior basolateral amygdalar nucleus — areas presumably modulating somatomotor outflow. Third, the BSTju sends dense projections to the caudal substantia innominata, a distinct caudal dorsolateral region of the compact part of the substantia nigra, and the adjacent mesencephalic reticular nucleus and retrorubral area. And fourth, the BSTju provides light inputs to the prelimbic, infralimbic, and ventral CA1 cortical areas; to the posterior basolateral, posterior basomedial, and lateral amygdalar nuclei; to the paraventricular and medial mediodorsal thalamic nuclei; to the subthalamic and parasubthalamic nuclei of the hypothalamus; and to the ventrolateral periaqueductal gray. These projections, in part, suggest a role for the BSTju in circuitry integrating autonomic responses with somatomotor activity in adaptive behaviors.     

Descending projections of the posterior nucleus of the hypothalamus: Phaseolus vulgaris leucoagglutinin analysis in the rat

No previous report in any species has systematically examined the descending projections of the posterior nucleus of the hypothalamus (PH). The present report describes the descending projections of the PH in the rat by using the anterograde anatomical tracer, Phaseolus vulgaris leucoagglutinin. PH fibers mainly descend to the brainstem through two routes: dorsally, within the central tegmental tract; and ventromedially, within the mammillo-tegmental tract and its caudal extension, ventral reticulo-tegmental tracts. PH fibers were found to distribute densely to several nuclei of the brainstem. They are (from rostral to caudal) 1) lateral/ventrolateral regions of the diencephalo-mesopontine periaqueductal gray (PAG); 2) the peripeduncular nucleus; 3) discrete nuclei of pontomesencephalic central gray (dorsal raphe nucleus, laterodorsal tegmental nucleus, and Barrington's nucleus); 4) the longitudinal extent of the central core of the mesencephalic through medullary reticular formation (RF); 5) the ventromedial medulla (nucleus gigantocellularis pars alpha, nucleus raphe magnus, and nucleus raphe pallidus); 6) the ventrolateral medulla (nucleus reticularis parvocellularis and the rostral ventrolateral medullary region); and 7) the inferior olivary nucleus. PH fibers originating from the caudal PH distribute much more heavily than those from the rostral PH to the lower brainstem. The PH has been linked to the control of several important functions, including respiration, cardiovascular activity, locomotion, antinociception, and arousal/wakefulness. It is likely that descending PH projections, particularly those to the PAG, the pontomesencephalic RF, Barrington's nucleus, and parts of the ventromedial and ventrolateral medulla, serve a role in a PH modulation of complex behaviors involving an integration of respiratory, visceromotor, and somatomotor activity. © 1996 Wiley-Liss, Inc.     

Anatomical and physiological characteristics of vestibular neurons mediating the vertical vestibulo-ocular reflexes of the squirrel monkey

The morphology of 35 vestibular neurons whose firing rate was related to vertical eye movements was studied by injection of horseradish peroxidase intracellularly into physiologically identified vestibular axons in alert squirrel monkeys. The intracellularly injected cells were readily classified into four main groups. One group of cells, down position-vestibular-pause neurons (down PVPs; N = 12), increased their firing rate during downward eye positions, paused during saccades, and were located in the medial vestibular nucleus (MV) and the adjacent ventrolateral vestibular nucleus (VLV). They had axons that crossed the midline and ascended in the medial longitudinal fasciculus (MLF) to terminate in the trochlear nucleus, the lateral aspect of the caudal oculomotor nucleus, and the dorsal aspect of the rostral oculomotor nucleus. A second group of cells (N = 15) were also located in the MV and VLV, but increased their firing rate during upward eye positions, and paused during saccades. These cells had axons that crossed the midline and ascended in the contralateral MLF to terminate in the medial aspect of the oculomotor nucleus. A third group of cells (N = 4) were located in the superior vestibular nucleus, generated bursts of spikes during upward saccades, and increased their tonic firing rate during upward eye positions. These cells had axons that ascended laterally to the ipsilateral MLF to terminate in regions of the trochlear and oculomotor nuclei similar to those in which down PVPs terminated. A fourth group of cells (N = 4), located in the VLV, had axons that projected to the spinal cord, although they had firing rates that were significantly correlated with vertical eye position. Electrical stimulation of the vestibular nerve evoked spikes at monosynaptic latencies in each of the above classes of cells, six of which were injected with horseradish peroxidase. Each group of cells had collateral projections to other areas of the brainstem. Some of the neurons that projected to the contralateral trochlear and oculomotor nuclei had collaterals that crossed the midline to terminate in the oculomotor nucleus ipsilateral to the soma, and some gave rise to small collaterals that terminated in the abducens nucleus. Other areas of the brainstem that received collateral inputs from neurons projecting to oculomotor and trochlear nuclei included the interstitial nucleus of Cajal, the caudal part of the dorsal raphe nucleus, the nucleus raphe obscurus, Roller's nucleus, the intermediate and caudal interstitial nuclei of the MLF, and the nucleus prepositus.     

Afferent projections to the caudal part of the dorsal nucleus of the raphe in cats

Horseradish peroxidase (HRP) was iontophoresed into the caudal part of the dorsal nucleus of the raphe (DNR) in cats. Labeled neurons with HRP were recognized in the medial and the superior vestibular nucleus. Electrical stimulation of the medial or the superior vestibular nucleus elicited orthodromic evoked potentials and unitary responses in the caudal part of the DNR. These projections may be involved in eye movement control. In addition, labeled neurons were located in the magnocellular division of the alaminar spinal trigeminal nucleus. This projection may be a part of the pathway conveying somatosensory inputs from the face to the cerebellar flocculus.     

Uncrossed and crossed projections from the upper cervical spinal cord to the cerebellar nuclei in the rat, studied by anterograde axonal tracing

In the upper cervical spinal segments, neurons in the medial part of lamina VI give rise to uncrossed spinocerebellar axons, whereas the central cervical nucleus (CCN) and neurons in laminae VII and VIII give rise to crossed spinocerebellar axons. Using anterograde labeling with biotinylated dextran in the rat, we examined the projections of these neuronal groups to the cerebellar nuclei. Uncrossed and crossed projections were distinguished by cerebellar lesions placed on the side contralateral or ipsilateral to the tracer injections confined to the second and third cervical spinal segments (C2 and C3, respectively). Labeled terminals of uncrossed projections were seen in the middle, dorsal, and ventrolateral parts of the middle subdivision and in the ventral part of the caudomedial subdivision of the medial nucleus. In the anterior interpositus nucleus, terminals were seen in the middle of the mediolateral extent, whereas, in the posterior interpositus nucleus, they were seen in lateral and caudal parts. The terminals of crossed projections from the CCN were distributed ventrally in medial to ventrolateral parts of the middle subdivision of the medial nucleus. Some terminals were seen in the caudomedial subdivision of the medial nucleus. In the anterior interpositus nucleus, labeled terminals were seen mainly in rostromedial parts, whereas, in the posterior interpositus nucleus, they were seen in caudal and dorsal parts of the medial half. The present study suggests that the medial lamina VI group and the CCN in the upper cervical segments project to the different areas of the cerebellar nuclei and are concerned with different functions.     

Connections of the caudal cerebellar interpositus complex in a new world monkey (Cebus apella)

The afferent and efferent connections of the cerebellar interpositus complex were studied in a capuchin monkey (Cebus apella) that had received a transcannular horseradish peroxidase implant into the caudal portion of the anterior interpositus nucleus and posterior interpositus nucleus. While the heaviest anterogradely labeled ascending projections were observed to the contralateral ventral posterolateral nucleus of the thalamus, pars oralis (VPLo), efferent projections were also observed to the contralateral ventrolateral thalamic nucleus (VLc) and central lateral (CL) nucleus of the thalamic intralaminar complex, magnocellular (and to a lesser extent parvicellular) red nucleus, nucleus of Darkschewitsch, zona incerta, nucleus of the posterior commissure, lateral intermediate layer and deep layer of the superior colliculus, dorsolateral periaqueductal gray, contralateral nucleus reticularis tegmenti pontis and basilar pontine nuclei (especially dorsal and peduncular), and dorsal (DAO) and medial (MAO) accessory olivary nuclei, ipsilateral lateral (external) cuneate nucleus (LCN) and lateral reticular nucleus (LRN), and to a lesser extent the caudal medial vestibular nucleus (MVN) and caudal nucleus prepositus hypoglossi (NPH), and dorsal medullary raphe. The heaviest retrograde labeling was corticonuclear Purkinje cells in the paramedian cerebellar cortex lateral to the vermis of lobules IV-VIII. Otherwise, retrogradely labeled sources of afferents were predominantly contralateral in the dorsal, dorsomedial, paramedian, and peduncular sectors of the basilar pons, NRTP, and dorsal accessory (DAO) and medial accessory (MAO) of olivary nuclei, but were predominantly ipsilateral in the LCN, LRN, and in the medullary reticular formation along the roots of the hypoglossal (XII) cranial nerve. It appeared that the connections with the contralateral dorsal basilar pons, NRTP, DAO and MAO, and ipsilateral LCN and LRN are reciprocal.(ABSTRACT TRUNCATED AT 250 WORDS)     

Topographic organization of subcortical projections to the anterior thalamic nuclei in the rat.

Subcortical projections to the anterior thalamic nuclei were studied in the rat, with special reference to projections from the mammillary nuclei, by retrograde and anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase. The medial mammillary nucleus (MM) projects predominantly ipsilaterally to the entire anterior thalamic nuclei, whereas the lateral mammillary nucleus projects bilaterally to the anterodorsal nucleus (AD) of the anterior thalamic nuclei. A topographic relationship was recognized between the MM and the anterior thalamic nuclei. The dorsal region of the pars mediana of the MM projects to the interanteromedial nucleus (IAM), whereas the ventral region projects to the rostral part of the anteromedial nucleus (AM). The dorsal and the ventral regions of the pars medialis project to the dorsomedial part of the AM at its caudal and rostral levels, respectively. The dorsomedial region of the pars lateralis projects to the ventral AM. The ventrolateral region of the pars lateralis projects to the ventral part of the anteroventral nucleus (AV) in such a manner that rostral cells project rostrally and caudal cells project caudally. The pars basalis projects predominantly ipsilaterally to the dorsolateral AV and bilaterally to the AD. The rostrolateral region of the pars posterior projects to the lateral AV, whereas the medial and the caudal regions of the pars posterior project to the dorsomedial AV. The rostrodorsal part of the nucleus reticularis thalami was found to project to the anterior thalamic nuclei; cells located rostrally in this part project to the IAM and AM, whereas cells located caudodorsally project to the AV and AD. The laterodorsal tegmental nucleus projects predominantly ipsilaterally to the AV, especially to its dorsolateral part. The present study demonstrates that subdivisions of the subcortical structures are connected to the subnuclei of the anterior thalamic nuclei, with a clear-cut topography arranged in the dorsoventral and the rostrocaudal dimensions.     

Subregions of the periaqueductal gray topographically innervate the rostral ventral medulla in the rat.

Previous anatomical and physiological studies have revealed a substantial projection from the periaqueductal gray (PAG) to the nucleus paragigantocellularis (PGi). In addition, physiological studies have indicated that the PAG is composed of functionally distinct subregions. However, projections from PAG subregions to PGi have not been comprehensively examined. In the present study, we sought to examine possible topographic specificity for projections from subregions of the PAG to PGi. Pressure or iontophoretic injections of wheat germ agglutinin-conjugated horseradish peroxidase, or of Fluoro-Gold, placed into the PGi of the rat retrogradely labeled a substantial number of neurons in the PAG from the level of the Edinger-Westphal nucleus to the caudal midbrain. Retrogradely labeled neurons were preferentially aggregated in distinct subregions of the PAG. Rostrally, at the level of the oculomotor nucleus, labeled neurons were i) compactly aggregated in the ventromedial portion of the PAG corresponding closely to the supraoculomotor nucleus of the central gray, ii) in the lateral and ventrolateral PAG, and iii) in medial dorsal PAG. More caudally, retrogradely labeled neurons became less numerous in the dorsomedial PAG but were more widely scattered throughout the lateral and ventrolateral parts of the PAG. Only few retrogradely labeled neurons were found in the ventromedial part of the PAG at caudal levels. Injections of retrograde tracers restricted to subregions of the PGi suggested topography for afferents from the PAG. Injections into the lateral portion of the PGi yielded the greatest number of labeled neurons within the rostral ventromedial PAG. Medially placed injections yielded numerous retrogradely labeled neurons in the lateral and ventrolateral PAG. Injections placed in the rostral pole of the PGi (medial to the facial nucleus) produced the greatest number of retrogradely labeled neurons in the dorsal PAG. To examine the pathways taken by fibers projecting from PAG neurons to the medulla, and to further specify the topography for the terminations of these afferents in the PGi, the anterograde tracer Phaseolus vulgaris-leucoagglutinin was iontophoretically deposited into subregions of the PAG that contained retrogradely labeled neurons in the above experiments. These results revealed distinct fiber pathways to the rostral medulla that arise from the dorsal, lateral/ventrolateral, and ventromedial parts of the PAG. These injections also showed that there are differential but overlapping innervation patterns within the PGi. Consistent with the retrograde tracing results, injections into the rostral ventromedial PAG near the supraoculomotor nucleus yielded anterograde labeling immediately ventral to the nucleus ambiguus in the ventrolateral medulla, within the retrofacial portion of the PGi.(ABSTRACT TRUNCATED AT 400 WORDS)     

Projections from the parabrachial nucleus to the vestibular nuclei: potential substrates for autonomic and limbic influences on vestibular responses

Previous anatomical studies in rabbits and rats have shown that the superior vestibular nucleus (SVN), medial vestibular nucleus (MVN) and inferior vestibular nucleus (IVN) project to the parabrachial nucleus (PBN) and K?lliker-Fuse (KF) nucleus. Adult male albino rabbits and Long-Evans rats received iontophoretic injections of biotinylated dextran amine, Phaseolus vulgaris leucoagglutinin, Fluoro-Gold or tetramethylrhodamine dextran amine into either the vestibular nuclei or the PBN and KF nuclei. The results were similar in both rats and rabbits. Injections of retrograde tracers into the vestibular nuclei produced retrogradely labeled neurons bilaterally in caudal third of the medial, external medial, and external lateral PBN in both species, with more variable labeling in KF. Rats also had consistent bilateral (predominantly contralateral) labeling in the ventrolateral PBN. The most prominent labeling was produced from injections that included the SVN, with fewer labeled neurons observed from injections in the caudal MVN and the IVN. Anterograde transport of BDA from injections into the PBN and KF nuclei of rabbits revealed prominent projections to the SVN, dorsal aspect of the rostral MVN, caudal MVN, pars beta of the LVN and IVN. These connections appear to contain a component that is reciprocal to the vestibulo-parabrachial pathway and a non-reciprocal component to regions connected with the vestibulocerebellum and vestibulo-motor reflex pathways. These connections support the concept that a synthesis of autonomic, vestibular and limbic information is an integral property of pathways related to balance control in both the brain stem and forebrain. It is suggested that these projections may contribute broadly to both performance tradeoffs in vestibular-related pathways during variations in the behavioral context and affective state and the close association between anxiety and balance function.     

Interpeduncular nucleus afferents in the rat

Afferents to the interpeduncular nucleus (IPN) of the rat were studied with the horseradish peroxidase (HRP) retrograde transport method. HRP was deposited microelectrophoretically in the IPN of adult rats. Major projections to the IPN originate in the medial habenular nucleus, the region surrounding the dorsal tegmental nucleus (accessory dorsal tegmental nucleus and the so-called dorsal tegmental nucleus pars lateralis), and the midbrain raphe (nucleus centralis superior and nucleus raphe dorsalis). Also, minor projections originate in the central gray and nucleus locus coeruleus. Our results indicate that the habenulointerpeduncular projection originates solely from the medial habenular nuclei and is topographically organized; medial regions of the medial habenular nuclei project to ventral portions of IPN and lateral regions project to the dorsal IPN.     

Interpeducular nucleus afferents in the rat

Afferents to the interpeduncular nucleus (IPN) of the rat were studied with the horseradish peroxide (HRP) retrograde transport method. HRP was deposited microelectrophoretically in the IPN of adult rats. Major projections to the IPN originate in the medial habenular nucleus, the region surrounding the dorsal tegmental nucleus (accessory dorsal tegmental nucleus and the so-called dorsal tegmental nucleus pars lateralis), and the midbrain raphe (nucleus centralis superior and nucleus raphe dorsalis). Also, minor projections originate in the central gray and nucleus locus coeruleus. Our results indicate that the habenulointerpeduncular projection originates solely from the medial habenular nuclei and is topographically organized; medial regions of the medial habenular nuclei project to ventral portions of IPN and lateral regions project to the dorsal IPN.     

Basic organization of projections from the oval and fusiform nuclei of the bed nuclei of the stria terminalis in adult rat brain.

The organization of axonal projections from the oval and fusiform nuclei of the bed nuclei of the stria terminalis (BST) was characterized with the Phaseolus vulgaris-leucoagglutinin (PHAL) anterograde tracing method in adult male rats. Within the BST, the oval nucleus (BSTov) projects very densely to the fusiform nucleus (BSTfu) and also innervates the caudal anterolateral area, anterodorsal area, rhomboid nucleus, and subcommissural zone. Outside the BST, its heaviest inputs are to the caudal substantia innominata and adjacent central amygdalar nucleus, retrorubral area, and lateral parabrachial nucleus. It generates moderate inputs to the caudal nucleus accumbens, parasubthalamic nucleus, and medial and ventrolateral divisions of the periaqueductal gray, and it sends a light input to the anterior parvicellular part of the hypothalamic paraventricular nucleus and nucleus of the solitary tract. The BSTfu displays a much more complex projection pattern. Within the BST, it densely innervates the anterodorsal area, dorsomedial nucleus, and caudal anterolateral area, and it moderately innervates the BSTov, subcommissural zone, and rhomboid nucleus. Outside the BST, the BSTfu provides dense inputs to the nucleus accumbens, caudal substantia innominata and central amygdalar nucleus, thalamic paraventricular nucleus, hypothalamic paraventricular and periventricular nuclei, hypothalamic dorsomedial nucleus, perifornical lateral hypothalamic area, and lateral tegmental nucleus. Moderately dense inputs are found in the parastrial, tuberal, dorsal raphé, and parabrachial nuclei and in the retrorubral area, ventrolateral division of the periaqueductal gray, and pontine central gray. Light projections end in the olfactory tubercle, lateral septal nucleus, posterior basolateral amygdalar nucleus, supramammillary nucleus, and nucleus of the solitary tract. These and other results suggest that the BSTov and BSTfu are basal telencephalic parts of a circuit that coordinates autonomic, neuroendocrine, and ingestive behavioral responses during stress.     

Dorsal mesencephalic projections to pons, medulla, and spinal cord in the cat: limbic and non-limbic components.

The vertebrate dorsal mesencephalon consists of the superior colliculus, the dorsal portion of the periaqueductal gray, and the mesencephalic trigeminal neurons in between. These structures, via their descending pathways, take part in various behavioral responses to environmental stimuli. This study was undertaken to compare the origins and trajectories of these pathways in the cat. Injections of horseradish peroxidase into the cervical spinal cord and upper medullary medial tegmentum retrogradely labeled cells mainly in the contralateral intermediate and deep superior colliculus, and in the ipsilateral dorsal and lateral periaqueductal gray and adjacent tegmentum. Only injections in the medullary lateral tegmental field labeled mesencephalic trigeminal neurons ipsilaterally. Autoradiographic tracing results, based on injections across the dorsal mesencephalon, revealed three efferent fiberstreams. A massive first fiberstream (limbic pathway), consisting of thin fibers, descended ipsilaterally from the dorsal and lateral periaqueductal gray and adjacent superior colliculus through the mesencephalic and pontine lateral tegmentum, terminating in these areas as well as in the ventral third of the caudal pontine and medullary medial tegmentum. A few fibers from the dorsal periaqueductal gray matter (PAG) were distributed bilaterally to the dorsal vagal, solitary, and retroambiguus nuclei. The second fiberstream (the predorsal bundle) descended contralaterally from the superior colliculus (SC) and consisted of both thick and thin labeled fibers. The thin fibers terminated bilaterally in the dorsomedial nucleus reticularis tegmenti pontis and the medial half of the caudal medial accessory inferior olive. The thick fibers targeted the contralateral dorsal two thirds of the caudal pontine and medullary medial tegmental fields, and the facial, abducens, lateral reticular, subtrigeminal, and prepositus hypoglossi nuclei. A few fibers recrossed the midline to terminate in the ipsilateral medial tegmentum. Caudal to the obex, fibers terminated laterally in the tegmentum and upper cervical intermediate zone. From the lateral SC, fibers terminated bilaterally in the lateral tegmental fields of the pons and medulla and lateral facial subnuclei. The third fiberstream (mesencephalic trigeminal or Probst tract) terminated in the supratrigeminal and motor trigeminal nuclei, and laterally in the tegmentum and upper cervical intermediate zone. In summary, neurons in the PAG and in the deep layers of the SC give rise to a massive ipsilateral descending pathway, in which a medial-to-lateral organization exists. A similar topographical pattern occurs in the crossed SC projections. The possibility that these completely different descending systems cooperate in producing specific defensive behaviors is discussed.     

The topographical organization of neurons in the rat medial frontal, insular and olfactory cortex projecting to the solitary nucleus, olfactory bulb, periaqueductal gray and superior colliculus

In 19 rats two different retrograde tracers (Fast Blue, Diamidino Yellow, Rhodamine-labeled latex microspheres, or wheat germ agglutinin conjugated with HRP) were injected into the solitary nucleus (NTS) and either the olfactory bulb (OB), periaqueductal gray (PAG) or superior colliculus (SC). The pattern of retrogradely labeled neurons in the medial frontal, insular and olfactory cortices was examined to determine the topographical organization of the cell populations projecting to these subcortical targets and the extent to which they overlapped. In the medial frontal cortex (MFC) SC projections originated most dorsally, while NTS and OB projections originated most ventrally and exhibited slight overlap. PAG projections originated from virtually the entire MFC and overlapped with cells projecting to the OB, NTS and SC. These results are consistent with the role of dorsal MFC as the rat's frontal eye field and the ventral MFC as a visceral motor area. Laterally, in the insular cortex there was virtually complete overlap between cells projecting to the NTS and PAG. The extensive overlap of PAG projections with NTS projections medially and laterally and with SC projections medially suggests the PAG is involved in a variety of brain visceral and somatic functions. In the piriform cortex there was overlap between cells projecting to the OB and cells projecting to the SC; the cells projecting to the SC were located in the endopiriform nucleus, and may provide a substrate for orienting responses to odors.     

Superior colliculus connections with visual thalamus in gray squirrels (Sciurus carolinensis): evidence for four subdivisions within the pulvinar complex

As diurnal rodents with a well-developed visual system, squirrels provide a useful comparison of visual system organization with other highly visual mammals such as tree shrews and primates. Here, we describe the projection pattern of gray squirrel superior colliculus (SC) with the large and well-differentiated pulvinar complex. Our anatomical results support the conclusion that the pulvinar complex of squirrels consists of four distinct nuclei. The caudal (C) nucleus, distinct in cytochrome oxidase (CO), acetylcholinesterase (AChE), and vesicular glutamate transporter-2 (VGluT2) preparations, received widespread projections from the ipsilateral SC, although a crude retinotopic organization was suggested. The caudal nucleus also received weaker projections from the contralateral SC. The caudal nucleus also projects back to the ipsilateral SC. Lateral (RLl) and medial (RLm) parts of the previously defined rostral lateral pulvinar (RL) were architectonically distinct, and each nucleus received its own retinotopic pattern of focused ipsilateral SC projections. The SC did not project to the rostral medial (RM) nucleus of the pulvinar. SC injections also revealed ipsilateral connections with the dorsal and ventral lateral geniculate nuclei, nuclei of the pretectum, and nucleus of the brachium of the inferior colliculus and bilateral connections with the parabigeminal nuclei. Comparisons with other rodents suggest that a variously named caudal nucleus, which relays visual inputs from the SC to temporal visual cortex, is common to all rodents and possibly most mammals. RM and RL divisions of the pulvinar complex also appear to have homologues in other rodents.     

Corticofugal connections between the cerebral cortex and the vestibular nuclei in the rat

The distribution of cortical efferent connections to the vestibular nuclei was quantitatively analyzed by means of retrograde axonal transport of horseradish peroxidase, wheat germ agglutinin-horseradish peroxidase, and Fast Blue in rats. The tracer substances were injected into the spinal vestibular nucleus (SpVe), the caudal part of the medial vestibular nucleus (MVe), and nucleus X of Brown Norwegian rats. Projections to the vestibular nuclei were revealed bilaterally, but predominantly contralaterally from five cortical areas: (1) the parietotemporal region (PT) which occupied the caudal two-thirds of the secondary somatosensory area and spread over the caudal part of the primary somatosensory area and the visceral cortex; (2) the anterior forelimb (AF) overlapping the anterior part of the forelimb area and the transitional zone; (3) the anterior hindlimb (AH) overlapping the anterior part of the hindlimb area and the transitional zone; (4) the lateral forelimb (LF) centered in the intercalated zone lateral to the forelimb area; and (5) the ventrotemporal region (VT) located at the ventral part of the temporal cortex. In addition to these cortical fields, the frontal cortex was found to project directly to the vestibular nuclei. These corticofugal projections were verified in experiments in which biocytin was injected into the rat PT. Anterogradely labelled fibers were traced predominantly contralaterally to the SpVe, caudal part of the MVe, and nucleus X. It is suggested that the rat corticofugal projections to the caudal vestibular nuclei modify vestibular reflexes to assist in coordinating eye, head and body movements during locomotion.