Spinal and trigeminal dorsal horn projections to the parabrachial nucleus in the rat

We studied afferents to the parabrachial nucleus (PB) from the spinal cord and the spinal trigeminal nucleus pars caudalis (SNVc) in the rat by using the anterograde and retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). Injections of WGA-HRP into medial PB retrogradely labeled neurons in the promontorium and in lamina I of the dorsal rostral SNVc, while injections into lateral PB and the K?lliker-Fuse nucleus retrogradely labeled neurons in these areas as well as in lamina I throughout the caudal SNVc and spinal dorsal horn. Injections of WGA-HRP into the caudal SNVc and dorsal horn of the spinal cord resulted in terminal labeling in the dorsal, central, and external lateral subnuclei of PB and the K?lliker-Fuse nucleus, all of which are known to receive cardiovascular and respiratory afferent information. Injections of WGA-HRP into the promontorium and dorsal rostral SNVc resulted in terminal labeling in the same PB subnuclei, as well as in the medial and the ventral lateral PB subnuclei, which are sites of relay for gustatory information ascending from the medulla to the forebrain. The spinal and trigeminal projection to PB may mediate the convergence of pain, chemosensory, and temperature sensibilities with gustatory and cardiorespiratory systems in PB.     

Cells of origin of the spinohypothalamic tract in the rat

We recently demonstrated that large numbers of neurons in the spinal cord of rats project directly to the hypothalamus. In the present study, we used the retrograde tracer Fluoro-Gold (FG) to examine this projection more completely. In the first series of studies, we attempted to label the entire population of spinal cord neurons that project to the hypothalamus. Injections that virtually filled the hypothalamus on one side without spreading into any other diencephalic area labeled a large number of neurons (estimated to be more than 9,000 in the case with the most neurons labeled) bilaterally at all levels of the spinal cord. Approximately 60% of the labeled neurons were contralateral to the injection. The greatest number of labeled neurons was found within the deep dorsal horn. Many were also found within the lateral spinal nucleus, the superficial dorsal horn, and the gray matter surrounding the central canal. A small number of labeled cells was located in the intermediate zone and ventral horn of the spinal gray matter. Labeled neurons were distributed bilaterally within the sacral parasympathetic nucleus and trigeminal nucleus caudalis. Injections of FG restricted to the medial hypothalamus labeled neurons within the spinal cord in a distribution similar to that produced by injections that filled the hypothalamus. However, fewer neurons were labeled in the spinal cord (estimated to be more than 6,200) and trigeminal nucleus caudalis. Injections of FG restricted to the lateral hypothalamus also labeled fewer neurons (approximately 3,300) than did injections that filled the hypothalamus. In these cases, also, the pattern of labeled neurons within the spinal cord was similar to that produced by injections within either medial or both medial and lateral hypothalamus. However, few neurons were labeled in the sacral parasympathetic nucleus following injections into the lateral hypothalamus. These findings show the distribution of a large number of spinal cord neurons that project directly to medial or lateral hypothalamic regions that are involved in autonomic, neuroendocrine, and emotional responses to somatosensory stimulation, including painful stimuli.     

Distinct lateral and medial projections of the spinohypothalamic tract of the rat

We recently described a direct nociceptive projection from the spinal cord to the hypothalamus in the rat. Several electrophysiological studies of this projection indicated that the axons of some spinohypothalamic tract neurons (SHT) reach the hypothalamus either by a lateral or by a medial route. The purpose of this study was to determine the origin of all SHT neurons that reach the hypothalamus through the lateral and the medial projections, and to investigate the possibility of ablating the SHT without damaging other important sensory and motor tracts by combining retrograde tracing techniques with axonal ablation. As compared with control cases, significant (P < .05) reductions in the number of labeled SHT neurons were encountered, 26% in the ipsilateral spinal cord following lesions of the medial projection, 67% in the contralateral spinal cord following lesions of the lateral projection, and 94% in both contra- and ipsilateral sides following lesions of both the medial and lateral projections. Bilateral lesions of the lateral projections had no effect on the distribution of labeled neurons in the spinal cord and dorsal column nuclei following injections of Fluoro-Gold (FG) into the thalamus, and a small unilateral lesion of the lateral projection reduced the ipsilateral labeling in the motor cortex following injections of FG into the pyramidal decussation. These findings suggest that most SHT neurons ascend through the contralateral lateral projection and that less than half continue in the medial projection to the ipsilateral side. They also suggest a site that can be lesioned without affecting other ascending sensory spinal pathways. © 1996 Wiley-Liss, Inc.     

Brainstem afferents to the intralaminar nuclei of the cat thalamus studied by the horseradish peroxidase method

Brainstem afferents to the intralaminar thalamic nuclei of the cat have been studied by retrograde axonal transport of horseradish peroxidase (HRP) or wheat germ agglutinin conjugated to HRP (WGA-HRP). Injections of HRP or WGA-HRP into the lateral central and paracentral nuclei led to labeling of cells in various diencephalic structures of the injected side. Some cells were labeled in the intralaminar and medial pulvinar nuclei. A large number of labeled cells were detected in the thalamic reticular and ventral lateral geniculate nuclei. The zona incerta and field of Forel contained a few labeled cells. Occasional labeled cells were seen in the dorsomedial, ventromedial, posterior and lateral hypothalamic nuclei. Labeling of cells in the more caudal brainstem structures was found bilaterally, mostly with ipsilateral predominance. Some labeled cells were detected in the pretectal nuclei, the periaductal gray and the pars reticulata of the substantia nigra as well. The nuclei of Darkschewitsch and Cajal contained occasional labeled cells. Numerous cells were labeled in the superior colliculus, mainly in layers 3 and 4. A number of labeled cells were distributed in almost the entire extent of the brainstem reticular formation. It should be mentioned that the gigantocellular reticular nucleus contained labeled cells bilaterally with contralateral predominance. Very few labeled cells were encountered in the ipsilateral dorsal tegmental and parabrachial nuclei. A few cells were labeled in the vestibular nuclei and nucleus prepositus hypoglossi bilaterally. Some labeled cells were detected mainly contralaterally in three cerebellar nuclei; the fastigial, the interposed and the dentate. Injections of WGA-HRP into the medial central nucleus showed a different pattern of labeling. It is noteworthy that no labeled cells were found in the dorsal thalamus, the hypothalamus (except for the lateral mamillary nucleus), the superior colliculus or the cerebellar nuclei.     

Afferent projections to the central nucleus of the inferior colliculus in the rat

The afferent projections to the inferior colliculus of the rat were studied using the method of retrograde transport of horseradish peroxidase (HRP).Following large injections of HRP into the central nucleus, cells within the cochlear nuclei, superior olivary complex and auditory cortex were stained. Within the contralateral dorsal cochlear nucleus, fusiform cells were heavily labeled. Giant cells were also labeled in deeper layers. In the contralateral ventral cochlear nucleus, virtually all major cell types were labeled, with some types being labeled in greater numbers than others. Octopus cells of posteroventral division of ventral cochlear nucleus (PVCN) were never labeled. HRP-positive cells were found in ipsilateral and contralateral lateral superior olivary nucleus (LSO), ipsilateral medial superior olivary nucleus (MSO), ipsilateral and contralateral lateral nucleus of the trapezoid body (LTB), ipsilateral ventral nucleus of the trapezoid body (VTB), and ipsilateral superior paraolivary nucleus (SPN). Pyramidal cells of layer V of auditory cortex were heavily labeled.Small injections of HRP into the central nucleus resulted in labeled cells within restricted regions of the cochlear nuclei, superior olivary complex and auditory cortex. Injections into dorsal regions of the central nucleus resulted in cells labeled in ventral regions of the dorsal and ventral cochlear nuclei, and in lateral regions of LSO. These regions contain neurons which are considered to have low best frequencies. Injections placed in more ventral regions of the central nucleus led to labeling of cells in more dorsal regions of the cochlear nuclei and more medial regions of LSO in agreement with the tonotopical progressions within these structures.     

Afferents to the entorhinal cortex of the rat studied by the method of retrograde transport of horseradish peroxidase

The afferents to the parahippocampal area of the rat were studied with retrograde transport of horseradish peroxidase injected into the medial entorhinal cortex, lateral entorhinal cortex, parasubiculum, presubiculum, or a large injection which stained all these structures as well as the ventral hippocampus. Control rats were injected with horseradish peroxidase into the overlying visual cortex. Labeled neurons in brains with injections into the medial entorhinal cortex and the adjacent parasubicular region were found in the ipsilateral and contralateral presubicular region, the medial septal nucleus, the thalamic nucleus reuniens, the dorsal part of the lateral nucleus of thalamus, the anterior periventricular nucleus of the thalamus, and the dorsal raphe nucleus. Brains with injections into the lateral entorhinal cortex yielded labeled neurons in the medial septal nucleus, nucleus reuniens, dorsal raphe nucleus, and nucleus locus ceruleus. Injections into the presubiculum resulted, in addition, in labeling of neurons in the lateral nucleus of the thalamus. Control injections aimed at the sensory cortex overlying the parahippocampal area yielded labeled neurons in the medial septal nucleus, the dorsal lateral geniculate nucleus, and the nucleus locus ceruleus.     

Mamillary body in the rat: topography and synaptology of projections from the subicular complex, prefrontal cortex, and midbrain tegmentum

The retrograde and anterograde transport of horseradish peroxidase conjugated to wheat germ agglutinin (WGA-HRP) has been used to trace afferent connections of the rat mamillary body (MB) at the light and electron microscopic levels. Injections of WGA-HRP into different parts of the MB resulted in heavy retrograde labeling in the subicular complex, medial prefrontal cortex, and dorsal and ventral tegmental nuclei. Injections of WGA-HRP into each of these brain regions, respectively, resulted in anterograde labeling with specific distributions and characteristic synaptic organizations in the MB. Projections from the rostrodorsal and caudoventral subiculum terminated in a topographically organized laminar fashion in the medial mamillary nucleus bilaterally, whereas afferent projections from the presubiculum and parasubiculum terminated only in the lateral mamillary nucleus. Labeled axon terminals which originated from the subicular complex were characterized by round vesicles and formed asymmetric synaptic junctions with small-diameter dendrites and dendritic spines in the medial and lateral mamillary nuclei. Projections from the prefrontal cortex originated mainly in the infralimbic area and to a lesser degree in the prelimbic and anterior cingulate areas. Injections of tracer into these brain regions gave rise to dense labeling of axon terminals in the medial mamillary nucleus, pars medianus, and in the anterior dorsomedial portion of the pars medialis. The labeled terminals were characterized by round vesicles and formed asymmetric synaptic junctions with small-diameter dendrites and dendritic spines. Projections from the dorsal tegmental nucleus terminated in the ipsilateral lateral mamillary nucleus, whereas afferent projections from the anterior and posterior subnuclei of the ventral tegmental nucleus terminated topographically in the medial mamillary nucleus. The ventral tegmental nucleus, pars anterior projected to the midline region of the medial nucleus and the dorsolateral and ventromedial subdivisions of the pars posterior projected to medial and lateral parts of the medial nucleus, respectively. In contrast to the synaptic morphology of subicular complex and medial prefrontal cortex axon terminals in the MB, labeled axon terminals in the MB which originated from the midbrain tegmentum were characterized by pleomorphic vesicles and formed symmetric synaptic junctions with neuronal somata and proximal dendrites as well as distal dendrites and dendritic spines.(ABSTRACT TRUNCATED AT 400 WORDS)     

The thalamo-cortical projection of the nucleus submedius in the cat

The cortical projection of the nucleus submedius (Sm) was studied in the cat with the autoradiographic and horseradish peroxidase (HRP) methods. The results indicate that Sm projects topographically on to layer 3 of a distinct agranular cortical field that occupies the posterolateral gyrus proreus, the adjacent fundus of the rhinal sulcus, and the postero-ventral portion of the medial wall of the presylvian sulcus. This cortical field is denoted the ventrolateral orbital cortex (VLO), consonant with previous nomenclature in the rat (Krettek and Price, '77a). The more ventral part (VLOβ) is cytoarchitectonically distinct from the dorsal part (VLOα); the former receives input from the anterior part of Sm (Sm_a), while the latter receives superficial layer 1 of VLO probably also arises from Sm, and there may be an input to layers 5 and 6. The corticothalamic projection from VLO to Sm reciprocates the ipsilateral thalamocortical projection and also has a moderate contralateral component. A dense, subpial layer 1 input to VLO arises from cells of the ventromedial nucleus (VM) subjacent to Sm. The present experiments also indicate that clusters of cells in VM probably provide input to layer 3 of the cortex in the fundus of the presylvian sulcus, as well as area 6aβ in the lateral wall of the presylvian sulcus and the ventral bank of the cruciate sulcus. Results from the HRP experiments additionally indicate that VLOβ and the anteroventral (Sm_v) portion of VLOα are reciprocally connected with the ventral agranular insular cortex and the cingulate cortex, ipsilaterally, while the posterodorsal (Sm_d) portion of VLOα is instead connected with specific portions of the somatosensory cortical areas bilaterally. All portions of VLOα appear to project to the ventrolateral periaqueductal gray. In light of the recent suggestion that (Sm_d) is involved with nociception (Craig and Burton, '81), the present results suggest that the related portion of VLOα may serve as a cortical representation for noxious stimuli.     

Direct cortical projections to the parabrachial nucleus in the cat

Direct projections from the cerebral cortex to the parabrachial nucleus in the cat were examined by the horseradish peroxidase (HRP)method. When HRP was injected into the parabrachial nucleus, retrogradely labeled neuronal cell bodies were seen, bilaterally with an ipsilateral predominance, mainly in the orbital gyrus, the lateral bank of the presylvian sulcus, and a restricted region in the infralimbic cortex on the medial surface of the frontal lobe (stereotaxic coordinates; Fr: 22, L: 1, H: -1); all labeled neurons were in deep pyramidal cell layer. After injecting HRP conjugated to wheat germ agglutinin (WGA-HRP) into the cortical regions where retrogradely labeled neurons were found after injecting HRP into the parabrachial nucleus, anterogradely labeled cortical fibers were traced to the parabrachial nucleus. Corticoparabrachial fibers originating from the orbital gyrus and the lateral bank of the presylvian sulcus ran ipsilaterally through the internal capsule and the cerebral peduncle down to the lower brainstem, whereas those from the infralimbic cortex coursed down ipsilaterally through the medial forebrain bundle. These cortical fibers to the parabrachial nucleus were distributed bilaterally with an ipsilateral predominance. Cortical fiber terminals in the parabrachial nucleus were topographically arranged: Corticoparabrachial fibers from the lateral bank of the presylvian sulcus ended most massively in the dorsal part of the lateral parabrachial nucleus. Corticoparabrachial fibers from the orbital gyrus ended most heavily in the medial parabrachial nucleus and less heavily in the lateral parabrachial nucleus. Corticoparabrachial fibers from the infralimbic cortex ended mostly in the parabrachial regions surrounding the brachium conjunctivum.     

Development of the projections from the dorsal lateral geniculate nucleus to the lateral suprasylvian visual area of cortex in the cat.

In the study reported in the preceding paper, we used retrograde labeling methods to show that the enhanced projection from the thalamus to the posteromedial lateral suprasylvian (PMLS) visual area of cortex that is present in adult cats following neonatal visual cortex damage arises at least partly from surviving neurons in the dorsal lateral geniculate nucleus (LGN). In the C layers of the LGN, many more cells than normal are retrogradely labeled by horseradish peroxidase (HRP) injected into PMLS cortex ipsilateral to a visual cortex lesion. In addition, retrogradely labeled cells are found in the A layers, which normally have no projection to PMLS cortex in adult cats. The purpose of the present study was to investigate the mechanisms of this enhanced projection by examining the normal development of projections from the thalamus, especially the LGN, to PMLS cortex. Injections of HRP were made into PMLS cortex on the day of birth or at 1, 2, 4, or 8 weeks of age. Retrogradely labeled neurons were present in the lateral posterior nucleus, posterior nucleus of Rioch, pulvinar, and medial interlaminar nucleus, as well as in the LGN, at all ages studied. Within the LGN of the youngest kittens, a small number of retrogradely labeled cells was present in the interlaminar zones and among the cells in the A layers that border these zones. Such labeled cells were virtually absent by 8 weeks of age, and they are not found in normal adult cats. Sparse retrograde labeling of C-layer neurons also was present in newborn kittens.(ABSTRACT TRUNCATED AT 250 WORDS)     

Thalamocortical and thalamo-amygdaloid projections from the parvicellular division of the posteromedial ventral nucleus in the cat

Projections from the parvicellular division of the posteromedial ventral thalamic nucleus (VPMpc) of the cat were examined. After injection of horseradish peroxidase conjugated with wheat germ agglutinin (WGA-HRP) into the VPMpc, both anterogradely labeled axon terminals and retrogradely labeled neuronal cell bodies were found ipsilaterally in three discrete regions of the cerebral cortex, i.e., in the orbital cortex, caudoventral part of the infralimbic cortex, and medial part of the fundus of the posterior rhinal sulcus (perirhinal area); in the subcortical regions, anterogradely labeled axon terminals were seen ipsilaterally in the rostrodorsal part of the lateral amygdaloid nucleus. Neuronal connections between these VPMpc-recipient regions were further verified by injecting WGA-HRP into each of the three cortical and the lateral amygdaloid regions. After injection of WGA-HRP into each of the three cortical regions, labeled neuronal cell bodies and axon terminals were seen ipsilaterally in the VPMpc, especially in its medial part, and in the other two of the three VPMpc-recipient cortical regions. In the rostrodorsal part of the lateral amygdaloid nucleus, both axon terminals and neuronal cell bodies were labeled after WGA-HRP injection into the perirhinal area, and only axon terminals were labeled after WGA-HRP injection into the orbital cortex, but no labeling was observed after WGA-HRP injection into the infralimbic cortex. After injection of WGA-HRP into the rostrodorsal portion of the lateral amygdaloid nucleus, both axon terminals and neuronal cell bodies were labeled ipsilaterally in the perirhinal area and the ectorhinal area, and only neuronal cell bodies were labeled ipsilaterally in the VPMpc (especially in its medial part) and orbital cortical region; no labeling was observed in the infralimbic cortex. The present results indicate that the VPMpc of the cat is connected reciprocally with the orbital, infralimbic, and perirhinal cortical regions on the ipsilateral side, that the three VPMpc-recipient cortical regions are reciprocally connected with each other, that the VPMpc sends fibers ipsilaterally to the rostrodorsal part of the lateral amygdaloid nucleus, which may relay information from the VPMpc to the perirhinal cortical area, and that the VPMpc-recipient area in the lateral amygdaloid nucleus receives cortical fibers from the orbital and perirhinal cortical regions.     

the connections of the mouse olfactory bulb: A study using orthograde and retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase

The efferent and centrifugal afferent connections of the main olfactory bulb (MOB) of the mouse were studied by orthograde and retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). MOB projects ipsilaterally to the anterior olfactory nucleus, taenia tecta, anterior hippocampal continuation, indusium grisium, olfactory tubercle, and the lateral and medial divisions of the entorhinal area. In the region of the anterior one-half to two-thirds of the posterior division of the insular cortex the projection from MOB extends into the insular cortex. The only efferent projection of MOB to the contralateral half of the brain was to the anterior olfactory nucleus. All efferent projections of MOB, thus, are to telencephalic structures. By contrast the centrifugal afferents to MOB originate from every major division of the neuraxis. Neurons projecting to the bulb were found ipsilaterally in all divisions of the anterior olfactory nucleus (AON). In some cases, labeling in the external division of AON was weak or absent. In the contralateral AON, pars externa was the most intensively labeled sub-division. Retrogradely labeled neurons were also present in all other subdivisions of the contralateral AON but were fewer in number and less heavily labeled than in the ipsilateral AON. Ipsilaterally, positive neurons were also present in taenia tecta, and the anterior hippocampal continuation. There was profuse retrograde labeling of neurons in the entire extent of the ipsilateral piriform cortex (PC). There was a rostral to caudal gradient of labeling in PC with more positive neurons in rostral than caudal parts. Labeled neurons were present in the lateral entorhinal cortex LEC and in the transitional cortex between LEC and PC. Very heavy retrograde labeling was present in the nuclei of the horizontal and vertical limbs of the diagonal band (HDB and VDB). More cells were labeled in HDB than in VDB. Neurons were labeled in the ipsilateral nucleus of the lateral olfactory tract (NLOT) and, when the injection spread into the accessory olfactory bulb, labeled neurons were present ventral to NLOT in accessory NLOT. A few lightly labeled neurons were always present in the posterolateral and medial cortical amygdaloid areas. Neurons were labeled in the zona inserta and scattered throughout several hypothalamic nuclei. There was massive retrograde labeling of neurons in the locus coeruleus and neurons were abundantly labeled in the dorsal and medial raphe nuclei and nucleus raphe pontis. In general, the labeling of MOB connections was more extensive than that which has been reported in closely related species.(ABSTRACT TRUNCATED AT 400 WORDS)     

Topography and synaptology of mamillary body projections to the mesencephalon and pons in the rat.

The anterograde and retrograde transport of horseradish peroxidase conjugated to wheat germ agglutinin (WGA-HRP) was used to study the anatomical organization of descending projections from the mamillary body (MB) to the mesencephalon and pons at light and electron microscopic levels. Injections of WGA-HRP into the medial mamillary nucleus resulted in dense anterograde and retrograde labeling in the ventral tegmental nucleus, while injections in the lateral mamillary nucleus resulted in dense anterograde labeling in the dorsal tegmental nucleus pars dorsalis and dense anterograde and retrograde labeling in the pars ventralis of the dorsal tegmental nucleus. Anterogradely labeled fibers in the mamillotegmental tract diverged from the principal mamillary tract in an extensive dorsocaudally oriented swath of axons which extended to the dorsal and ventral tegmental nuclei, and numerous axons turned sharply ventrally and rostrally to terminate topographically in the dorsomedial nucleus reticularis tegmenti pontis and rostromedial pontine nuclei. The anterograde labeling in these two precerebellar relay nuclei was distributed near the midline such that projections from the lateral mamillary nucleus terminated mainly dorsomedial to the terminal fields of projections from the medial mamillary nucleus. In the dorsal and ventral tegmental nuclei, labeled axon terminals contained round synaptic vesicles and formed asymmetric synaptic junctions primarily with small diameter dendrites and to a lesser extent with neuronal somata. A few labeled terminals contained pleomorphic vesicles and formed symmetric synaptic junctions with dendrites and neuronal somata. Labeled axon terminals were also frequently found in synaptic contact with retrogradely labeled dendrites and neuronal somata in the dorsal and ventral tegmental nuclei. These findings indicate that neurons in the dorsal and ventral tegmental nuclei are reciprocally connected with MB projection neurons. In the nucleus reticularis tegmenti pontis and medial pontine nuclei, labeled axon terminals contained round synaptic vesicles and formed asymmetric synaptic junctions primarily with small diameter dendrites. The present study demonstrates that projections from the medial and lateral nuclei of the MB are topographically organized in the mesencephalon and pons. The synaptic morphology of mamillotegmental projections suggests that they may have excitatory influences primarily on the distal dendrites of neurons in these brain regions.     

Topographic organization of medial pulvinar connections with the prefrontal cortex in the rhesus monkey

The medial nucleus of the pulvinar complex (PM) has widespread connections with association cortex. We investigated the connections of the PM with the prefrontal cortex (PFC) in macaque monkeys, with tracers placed into the PM and the PFC, respectively. Injections of wheat germ agglutinin-horseradish peroxidase (WGA-HRP) placed into the PM resulted in widespread anterograde terminal labeling in layers III and IV, and retrograde cellular labeling in layer VI of the PFC. Injections of tracers centered on the central/lateral PM resulted in labeling of dorsolateral and orbital regions, whereas injections centered on caudal, medial PM resulted in labeling of dorsomedial and medial PFC. Since injections of the PM included neighboring thalamic nuclei, retrograde tracers were placed into distinct cytoarchitectonic regions of the PFC and retrogradely labeled cells in the posterior thalamus were charted. The results of this series of tracer injections confirmed the results of the thalamic injections. Injections placed into areas 8a, 12 (lateral and orbital), 45, 46 and 11, retrogradely labeled neurons in the central/lateral PM, while tracer injections placed into areas 9, 12 (lateral), 10 and 24, labeled medial PM. The connections of the PM with temporal, parietal, insular, and cingulate cortices were also examined. The central/lateral PM has reciprocal connections with posterior parietal areas 7a, 7ip, and 7b, insular cortex, caudal superior temporal sulcus (STS), caudal superior temporal gyrus (STG), and posterior cingulate, whereas medial PM is connected mainly with the anterior STS and STG, as well as the cingulate cortex and the amygdala. These connectional studies suggest that the central/lateral and medial PM have divergent connections which may be the substrate for distinct functional circuits. J. Comp. Neurol. 379:313–332, 1997. © 1997 Wiley-Liss, Inc.     

Thalamic projections to the hippocampal and entorhinal areas in the cat

The thalamic projections to the hippocampal formation and to the subicular and entorhinal areas in the cat have been studied with retrograde transport of horseradish peroxidase (HRP) or wheat germ agglutinin conjugated to HRP (WGA-HRP) and anterograde transport of WGA-HRP. Retrograde transport tracers injected in various parts of these cortices resulted in labeled cells in the midline, anterior, and lateral dorsal nuclei. Injections into the hippocampal formation or the subiculum led to retrograde labeling of cells in the reuniens nucleus of the ipsilateral thalamus throughout its rostrocaudal extent, whereas the restricted injections into the dentate gyrus and the inferior region of the hippocampus led to no labeling. Following an injection into the pre- and parasubiculum, a large number of labeled cells were seen not only in the reuniens nucleus but in other midline nuclei. In addition, a substantial number of labeled cells were also detected in the anterior and lateral dorsal nuclei, particularly in the anterodorsal nucleus, which contained densely arranged labeled cells throughout almost the entire rostrocaudal extent. An injection into the medial entorhinal area labeled a number of cells in the anterior nuclei and in the reuniens nucleus, particularly its dorsal part. Injections into various subdivisions of the lateral entorhinal area yielded different patterns of distribution of labeled cells in the thalamic nuclei. An injection into the ventromedial division (VMEA) led to abundant labeling of cells in the paraventricular and reuniens nuclei. After an injection into the ventral division (VLEA), numerous labeled cells were detected in the reuniens nucleus and a lesser number in the paraventricular nucleus at anterior levels. When an injection was made into the dorsal division (DLEA), a large number of labeled cells were detected in the reuniens nucleus, and less numerous labeled cells were found in the central medial nucleus. There appears to be a topographic arrangement of cortical projections of the reuniens nucleus. The pre- and parasubiculum receive projections from the most medial part of the reuniens nucleus near the midline, and the DLEA receives projections from the medial part of the nucleus. The cells projecting to the VLEA and MEA are distributed in the central part of the reuniens nucleus, and those to the VMEA are distributed in the lateral part. Anterograde experiments were also performed; injections of WGA-HRP into the reuniens nucleus resulted in terminal labeling in the superficial layers of the subicular area and the neighboring hippocampus and in the entorhinal area.     

Afferent and efferent connections of the olfactory bulbs in the lizard Podarcis hispanica.

The connections of the olfactory bulbs of Podarcis hispanica were studied by tract-tracing of injected horseradish peroxidase. Restricted injections into the main olfactory bulb (MOB) resulted in bilateral terminallike labeling in the medial part of the anterior olfactory nucleus (AON) and in the rostral septum, lateral cortex, nucleus of the lateral olfactory tract, and ventrolateral amygdaloid nucleus. Bilateral retrograde labeling was found in the rostral lateral cortex and in the medial and dorsolateral AON. Ipsilaterally the dorsal cortex, nucleus of the diagonal band, lateral preoptic area, and dorsolateral amygdala showed labeled cell bodies. Retrogradely labeled cells were also found in the midbrain raphe nucleus. Results from injections into the rostral lateral cortex and lateral olfactory tract indicate that the mitral cells are the origin of the centripetal projections of the MOB. Injections in the accessory olfactory bulb (AOB) produced ipsilateral terminallike labeling of the ventral AON, bed nucleus of the accessory olfactory tract, central and ventromedial amygdaloid nuclei, medial part of the bed nucleus of the stria terminalis, and nucleus sphericus. Retrograde labeling of neurons was observed ipsilaterally in the bed nucleus of the accessory olfactory tract and stria terminalis, in the central amygdaloid nucleus, dorsal cortex, and nucleus of the diagonal band. Bilateral labeling of somata was found in the ventral AON, the nucleus sphericus (hilus), and in the mesencephalic raphe nucleus and locus coeruleus. Injections into the dorsal amygdala showed that the mitral neurons are the cells of origin of the AOB centripetal projections. Reciprocal connections are present between AOB and MOB. To our knowledge, this is the first study to address the afferent connections of the olfactory bulbs in a reptile. On the basis of the available data, a discussion is provided of the similarities and differences between the reptilian and mammalian olfactory systems, as well as of the possible functional role of the main olfactory connections in reptiles.     

Projections to the subcortical forebrain from anatomically defined regions of the medial geniculate body in the rat

Although the auditory cortex is believed to be the principal efferent target of the medial geniculate body (MG), our recent behavioral studies indicate that in rats the conditioned coupling of emotional responses to an acoustic stimulus is mediated by subcortical projections of the MG. In the present study we have therefore used WGA-HRP as an anterograde and retrograde axonal marker to (1) define the full range of subcortical efferent projections of the MG; (2) identify the cells of origin within the MG of each projection; and (3) determine whether the subregions of the MG that project to subcortical areas receive inputs from acoustic relay nuclei of the mid-brain, particularly the inferior colliculus. The rat MG was first parcelled into three major cytoarchitectural areas: the ventral, medial, and dorsal divisions. The suprageniculate nucleus, located within the body of the MG just dorsal to the medial division, was also identified. Efferent projections of the MG were determined by combined anterograde and retrograde tracing methods. Injections of WGA-HRP in the MG produced anterograde transport to cortex and several subcortical areas, including the posterior caudate-putamen and amygdala, the ventromedial nucleus of the hypothalamus, and the subparafascicular thalamic nucleus. The cells of origin of the subcortical projections were then mapped retrogradely after injections in the anterogradely labeled areas. Injections in the caudate-putamen or amygdala retrogradely labeled the medial division of the MG and the suprageniculate nucleus, as well as several adjacent areas of the posterior thalamus surrounding the MG. In contrast, injections in the ventromedial nucleus of the hypothalamus or the subparafascicular thalamic nucleus only produced labeling in the areas surrounding MG. Afferents to MG from the inferior colliculus were then identified. The central nucleus of the inferior colliculus, the main lemniscal acoustic relay nucleus in the midbrain, was found to project to the ventral and medial divisions of the MG. In contrast, the dorsal cortex and external nucleus of the inferior colliculus project to each division of the MG and to several additional nuclei in adjacent areas of the posterior thalamus. These data demonstrate that the medial division of MG, the suprageniculate nucleus, and immediately adjacent areas of the posterior thalamus provide a direct linkage between auditory neurons in the inferior colliculus and subcortical areas of the forebrain and thereby support the view that thalamic sensory nuclei relay afferent signals to subcortical as well as cortical areas.     

The inferior olive of Saimiri sciureus: Olivocerebellar projections to the anterior lobe

Olivocerebellar projections to lobules IV and V of the anterior lobe of the squirrel monkey (Saimiri sciureus) cerebellum were studied using a wheat germ agglutinin-horseradish peroxidase conjugate (WGA-HRP). Placements of WGA-HRP in the cerebellar cortex, labeled neurons in different subgroups of the contralateral inferior olive depending on the position of the injection site. Injections in medial aspects of lobules IV and V labeled cells in subgroups a and b of the caudal medial accessory olive (MAO) and in the lateral dorsal accessory olive. Rostral parts of subgroup a of MAO and medial dorsal accessory olive contained labeled neurons when injections were made in intermediate areas of the cerebellar cortex. Subsequent to involvement of more lateral parts of lobules IV and V, labeled cells were found in the dorsal lamella and lateral bend of the principal olive. When the position and size of injection sites were compared to distribution of retrogradely labeled olivary somata, it was clear that zones A, B, C1, C2, C3 and D were present in lobules IV and V of squirrel monkey. The general pattern of olivary labeling in this study is similar to that reported for other species.