Cortical and brain stem afferents to the ventral thalamic nuclei of the cat demonstrated by retrograde axonal transport of horseradish peroxidase

After horseradish peroxidase (HRP) injections into various parts of the ventral thalamic nuclear group and its adjacent areas, the distribution of labeled neurons was compared in the cerebral cortex, basal ganglia, and the brain stem. The major differences in distribution patterns were as follows: Injections of HRP into the lateral or ventrolateral portions of the ventroanterior and ventrolateral nuclear complex of the thalamus (VA-VL) produced retrogradely labeled neurons consistently in area 4 gamma (lateral part of the anterior and posterior sigmoid gyri, lateral sigmoid gyrus and the lateral fundus of the cruciate sulcus), the medial division of posterior thalamic group (POm), suprageniculate nucleus (SG) and anterior pretectal nucleus ipsilaterally, and in the nucleus Z of the vestibular nuclear complex bilaterally. Injections into the medial or dorsomedial portion of the VA-VL resulted in labeled neurons within the areas 6a beta (medial part of the anterior sigmoid gyrus), 6a delta (anterior part of ventral bank of buried cruciate sulcus), 6 if. fu (posterior part of the bank), fundus of the presylvian sulcus (area 6a beta), medial part of the nucleus lateralis posterior of thalamus and nucleus centralis dorsalis ipsilaterally, and in the entopeduncular nucleus (EPN) and medial pretectal nucleus bilaterally. Only a few neurons were present in the contralateral area 6a delta. After HRP injections into the ventral medial nucleus (VM), major labeled neurons were observed in the gyrus proreus, area 6a beta (mainly in the medial bank of the presylvian sulcus), and EPN ipsilaterally, and in the medial pretectal nucleus and substantia nigra bilaterally. Following HRP injections into the centre médian nucleus (CM), major labeled neurons were found in the areas 4 gamma, 6a beta, and the orbital gyrus ipsilaterally, and in the EPN, rostral and rostrolateral parts of the thalamic reticular nucleus, locus ceruleus, nucleus reticularis pontis oralis et caudalis and nucleus prepositus hypoglossi bilaterally. The contralateral intercalatus nucleus also possessed labeled neurons. With HRP injections into the paracentral and centrolateral nuclei, labeled neurons were observed in the gyrus proreus and the cortical areas between the caudal presylvian sulcus and anterior rhinal sulcus ipsilaterally, and in the nuclei interstitialis and Darkschewitsch bilaterally. Minor differences in the distribution pattern were observed in the superior colliculus, periaqueductal gray, mesencephalic and medullary reticular formations, and vestibular nuclei in all cases of injections.     

The identification of brainstem neurones projecting to thoracic respiratory motoneurones in the cat as demonstrated by retrograde transport of HRP

Brainstem neurones which project to the immediate vicinity of the spinal motoneurones which supply the intercostal and abdominal respiratory muscles were identified by means of the retrograde transport of horseradish peroxidase (HRP). A combined electrophysiological and histological technique was used in which recording of phasic inspiratory or expiratory motoneurone activity within upper (T3-T4) or lower (T8-T9) thoracic segments was followed by the ion-tophoretic injection of HRP at these recording sites. HRP labelled cells were concentrated in those brainstem regions known to contain phasic respiratory neurones, namely the ventrolateral nucleus of the solitary tract (vl-NTS) or dorsal respiratory group (DRG), the ambiguus complex or ventral respiratory group (VRG) and the parabrachial pontine (PB) nuclei. In 18 cats, 248 cells were labelled in these three respiratory regions of the brainstem while 668 were much more diffusely distributed in other regions of the medulla and pons. The ipsilateral and contralateral contributions within the respiratory regions were respectively; 23%:77% (DRG), 33%:67% (VRG), 95%:5% (PB). These results are considered in the general context of previous electrophysiological and histological findings, but also with particular reference to a related study of the projections from brainstem neurones to the phrenic nucleus [32].     

Anterograde transsynaptic transport of WGA-HRP from spinal afferents to postganglionic sympathetic cells of the stellate ganglion of the guinea pig.

After injection of wheat germ agglutinin conjugated horseradish peroxidase (WGA-HRP) or choleragenoid conjugated HRP (B-HRP) into lower cervical and upper thoracic dorsal root ganglia (DRG), HRP reaction product was observed in peripheral fibers of spinal afferents and in postganglionic cell bodies of the stellate ganglion (SG) in the guinea pig. After WGA-HRP injection into C8-T3 or T5 DRG, HRP-labelled cells were observed to cluster at the rami within the SG, with peak labelling observed 36 h after injection. SG cell labelling occurred with B-HRP as well, but not with native HRP after injection into thoracic DRG. Injection of this tracer in C8 DRG gave rise to a small number of labelled cells. In contrast to the labelling pattern following thoracic or C8 DRG injections, injection of WGA-HRP or native HRP into C6 DRG, led to random SG cell labelling. We conclude that the anterograde transsynaptic transport, following injection of WGA-HRP into thoracic DRG, provides a method to selectively label a population of postganglionic sympathetic neurons within the SG. A combination of transsynaptic and retrograde transport appears to be responsible for labelling after injection into C8 DRG, whereas labelling after C6 DRG injections seems to be due primarily to retrograde transport.     

Distribution of calcitonin gene-related peptide-like immunoreactivity in the nucleus ambiguus of the cat

The distribution of calcitonin gene-related peptide (CGRP) in the cat nucleus ambiguus was examined by means of a combination of horseradish peroxidase (HRP) tracing and immunohistochemical techniques. Vagal motoneurones in the nucleus ambiguus were identified by applying HRP to either the thoracic vagus or the superior laryngeal nerve or the cervical vagus. Motoneurones in the nucleus ambiguus labelled with HRP from the thoracic vagus did not contain CGRP-like immunoreactivity although CGRP-like immunoreactive cells were present in this nucleus on the same sections. In contrast, a large proportion of the motoneurones labelled from the superior laryngeal nerve and a smaller proportion of cells labelled from the cervical vagus did contain CGRP-like immunoreactivity. It is concluded that CGRP-like immunoreactivity is absent from vagal preganglionic motoneurones projecting to structures in the thorax and abdomen but is present in vagal motoneurones projecting to striated muscle of the larynx and pharynx.     

Sources of thalamic afferent neurons, projecting into the suprasylvian gyrus of the dolphin cerebral cortex

The sources of a thalamic input to different loci of the suprasylvian gyrus (SSG) of the porpoise (Phocaena phocaena) cortex were studied by means of the retrograde HRP and fluorescent tracing methods. After injections of HRP into the anterior part of the SSG most cells were labelled in the lateral part of the ventrobasal complex. Some cells were also labelled in the ventroposteroinferior nucleus, posterior nucleus and caudally in the ventral parvocellular medial geniculate (MG). After injections of bisbenzimide in the middle part of the SSG many labelled cells were found in the ventral parvocellular MG and in the inferior pulvinar. Less cells were labelled in the magnocellular MG, lateral pulvinar and posterior nucleus. After bisbenzimide injection into the posterior part of the SSG the similar distribution of labelled cells was found but a sheet of labelled cells was shifted more laterally.     

Cerebral cortical projections to the reticular regions around the trigeminal motor nucleus in the cat

Cerebral cortical regions which send projection fibers to the reticular regions around the trigeminal motor nucleus were identified in the cat by the horseradish peroxidase (HRP) method. The reticular region around the trigeminal motor nucleus are known to contain many interneurons for masticatory motoneurons. After injections of HRP into the reticular regions around the trigeminal motor nucleus, HRP-labeled neuronal cell bodies in the cerebral cortex were found in layer V. They were distributed bilaterally in the orbitofrontal cortical regions, mainly in the rostral extension of the orbital gyrus close to the presylvian sulcus; more were located in the floor and lateral bank of the presylvian sulcus than in the crown of the orbital gyrus. After injections of HRP conjugated with wheat germ agglutinin (WGA-HRP) into these cortical regions, many labeled presumed axon terminals were distributed bilaterally in the reticular regions around the trigeminal motor nucleus; mainly in the region ventral to the trigeminal motor nucleus and in the intertrigeminal region between the main sensory trigeminal nucleus and the trigeminal motor nucleus. Terminal labeling in these regions was more prominent after WGA-HRP injection into the lateral bank of the presylvian sulcus than after WGA-HRP injection into the crown of the orbital gyrus. Thus, the present results indicate that the main part of the cortical region projecting directly to the reticular regions around the trigeminal motor nucleus in the cat is folded into the presylvian sulcus.     

The thalamic connectivity of the primary motor cortex (MI) in the raccoon

The purpose of this study was to determine the pattern of thalamic projections of the primary motor cortex (MI) in the raccoon, a carnivore species noted for neural specialization of sensorimotor function. Following electrophysiological identification of circumscribed regions of MI, injections of horseradish peroxidase (HRP) or HRP combined with tritiated amino acids were made in 15 animals. Labeled thalamic cells were found predominantly in the ventral lateral nucleus (VL). For a given cortical injection site within MI, labeled neurons in VL formed a crescent-shaped band which extended in a dorsoventral direction when viewed in coronal sections. These bands were topographically organized. Following an injection of the MI hindlimb area in the medial part of the posterior cruciate gyrus, both retrogradely labeled neurons and anterograde label formed a thin band at the lateral edge of VL while an injection of the MI face area in the lateral part of the anterior cruciate gyrus resulted in both anterograde and retrograde label in medial VL and the principal division of the ventromedial nucleus (VMp). An injection of the MI forepaw area localized to the rostral and central part of the posterior cruciate gyrus resulted in a band of labeled neurons occupying the dorsal extent of VL and continuing into the ventrolateral aspect of the ventral anterior nucleus (VA). In contrast, an injection of the MI forepaw area which was localized to the caudal extent of the posterior cruciate gyrus resulted in a wide and diffuse band of labeled neurons and anterograde label in the ventral portion of VL. All injections of MI produced cell labeling in the paracentral nucleus (PC) and the central lateral nucleus (CL) of the intralaminar group. These results demonstrate that VL is the primary thalamic dependency of MI in the raccoon. Labeled cells were not observed in the ventrobasal complex. The MI pattern of thalamic connectivity observed in the present study suggests that while differences exist in the regional specialization of sensorimotor structures among species, there appears to be little variation in the overall organization of thalamocortical relations.     

The cells of origin of the primate spinothalamic tract.

Spinothalamic tract cells in the lumbar, sacral and caudal segments of the primate spinal cord were labelled by the retrograde transport of horseradish peroxidase (HRP) injected into the thalamus. The laminar distribution of stained spinothalamic cells in the lumbosacral enlargement differed according to whether the HRP was injected into the lateral or the medial thalamus. Lateral injections labelled cells in most laminae, but the largest numbers of cells were in laminae I and V. The highest concentrations of cells labelled from the medial thalamus were in laminae VI-VIII. Ninety percent or more of the stained spinothalamic cells in the lumbosacral enlargement were contralateral to the injection site. In the conus medullaris stained spinothalamic cells were most numerous in laminae I, V and VI following lateral thalamic injections of HRP. Many of the cells of the conus were in Stilling's nucleus. Twenty-three percent of the cells in the conus were ipsilateral to the injection site in the lateral thalamus. Only a few cells in the conus were labelled by medial thalamic injections. The total number of spinothalamic cells from L5 caudally was estimated to be at least 1,200-2,500. An injection of HRP into the midbrain resulted in laminar distribution of labelled cells much like that produced by a lateral thalamic injection. The types of spinothalamic tract cells and the sizes of their somata were determined for different laminae. The cell types resemble those already described from Golgi and other studies of the spinal cord gray matter. The spinothalamic tract cells in lamina I included Waldeyer cells and numerous small fusiform, pyriform or triangular cells. Those in lamina II included limitrophe and central cells. Spinothalamic cells in lamina III were central cells. Most of the labelled cells in laminae IV-X were polygonal, although there were also flattened cells in these layers. The smallest spinothalamic cells were in laminae I-III, while the largest were in laminae V and VII-IX. Spinothalamic cells in the conus medullaris included cells like those in the lumbosacral enlargement, but also a special cell type in Stilling's nucleus. Some cells in the conus had dendrites that crossed the midline. Spinothalamic axons could sometimes be traced to the ventral white commissure within one or a few sections. In longitudinal sections, most labelled axons were in the ventral part of the lateral funiculus on the side of the injection, although a few were in the ventral funiculus or on the contralateral side. The axons were widely dispersed, and a few were located adjacent to the pia-glial membrane.     

Raccoon forelimb motorsensory cortex: I. Somatic afferent inputs to different cytoarchitectonic areas

The distribution of potentials evoked in and around forelimb MsI by graded electrical stimulation of forelimb nerves has been studied in the raccoon (Procyon lotor). These data have been correlated with cytoarchitectonic characteristics of pericruciate cortical tissue. Potentials evoked by cutaneous nerve stimulation were widely distributed in MsI and SmI, but were smaller in amplitude and of longer latency in MsI than in SmI. Stimulation of ulnar, median or deep radial nerve at 1-1.4T, a strength considered to activate only Group I muscle afferent fibers, caused evoked potentials in a localized region mostly confined to posterior sigmoid gyrus. On the basis of cytoarchitectonic features it is concluded that: a) Anterior sigmoid gyrus, to near the level of the tip of the cruciate sulcus, is area 6 cortex; b) The lateral portion of the posterior sigmoid gyrus, cortex comprising the caudal bank of the cruciate sulcus and cortex surrounding the lateral tip of the cruciate sulcus is area 4 cortex; c) The middle portion of the posterior sigmoid gyrus, almost to the lip of the cruciate sulcus rostrally and extending onto the rostral bank of the ascending coronal and postcruciate sulci caudally, is area 3a cortex. The cortical focus for Group I afferent-evoked potentials coincides with area 3a cortex. It is concluded that forelimb MsI of raccoon is organized in a fashion similar to MsI of cats and monkeys.     

Characterization of the pigeon isthmo-tectal pathway by selective uptake and retrograde movement of radioactive compounds and by Golgi-like horseradish peroxidase labeling.

Horseradish peroxidase (HRP) injected into developing limb buds of Xenopus laevis tadpoles is carried by retrograde axonal transport to the somata of motoneurones in the ventral horn. Small injection of 10% HRP were found to remain well localised to specified sites in the limb bud. Two types of labelled cells were found: diffusely labelled and granular labelled. Diffusely labelled cells result from axonal damage in the presence of HRP. Granular labelled cells result only from uptake of HRP from the region of the axon endings. No gradular uptake was found from axon shafts. It is concluded that the distribution of granular labelled cells accurately reflects the region of the ventral horn projecting to the site of injection in the limb.     

Quantitative synaptology of feline motoneurones to external anal sphincter muscle

Motoneurones innervating the cat external anal sphincter muscle were labelled retrogradely following intramuscular injections with horseradish peroxidase (HRP). Labelled motoneurones were examined by correlative light and electron microscopy (LM and EM) with special regard to a qualitative and morphometric analysis of the axon terminals resident on the neuronal membrane. By LM, labelled motoneurones were (1) ipsilateral to the injections; (2) all in S1-S2; (3) found only in the superior dorsomedial region of Onuf's nucleus; and (4) exhibited a broad spectrum of diameters (25-72 micron, mean 47.4 +/- 11.3 micron). By EM, axon terminals on the neuronal membrane when classified according to size, vesicle shape, and synaptic complex ultrastructure conformed to the S-, F-, T-, M-, and C-type terminals previously described for cat lumbosacral motoneurones. C-terminals confirmed these sphincteric motoneurones to be skeletomotor. Pooled data from midnuclear sections through 15 random labelled motoneurones (20-64-micron diameter) revealed that S- and F-type terminals predominated, with numerically few M and C types. Notwithstanding their low frequency (0.3/100 micron membrane) C-terminals contributed 1% of the mean areal coverage by terminals, which implies a potentially larger synaptic influence relative to other terminal types. Linear relationships occurred between terminal frequency (or cover) and motoneurone diameter. While motoneurones greater than 40 micron in diameter exhibited all five terminal types, labelled motoneurones less than or equal to 30 micron generally possessed only S-, F-, and occasional T-type terminals, and in this respect resembled gamma motoneurones.     

Absence of postnatal death among motoneurones supplying the inferior gluteal nerve of the rat

Motoneurones innervating the caudal part of the gluteus maximus muscle of 0-2 day, 10-12 day and 2-3-month-old rats were labelled by a half-hour application of a solution of 30% horseradish peroxidase (HRP) and 2% lysophatidylcholine delivered by suction electrode to the cut inferior gluteal nerves. The numbers of motoneurones labelled 24-48 h later were not significantly different in the 3 age groups (mean = 58.75, 54.0, 57.5, respectively). When a simple 30% HRP solution was used in adult rats, the number of motoneurones labelled was significantly less (mean = 48.75). In contrast, application of 0.5 microliter of HRP in a pledget of gelfoam to either the cut or uncut inferior gluteal nerve of neonates labelled large numbers of motoneurones, presumably by diffusion into nearby muscles. It is concluded that no death of motoneurones innervating the gluteus maximus muscle occurs postnatally, and that spread of HRP to neighbouring muscles can give rise to spuriously high motoneurone counts in neonates, and that incomplete uptake or transport of HRP in adults can lead to incorrectly low counts.     

Inputs from motor and premotor cortex to the superior colliculus of the macaque monkey

In retrograde studies of corticotectal projections in the monkey using horseradish peroxidase (HRP), projections of the frontal lobes were found to originate not only from the frontal eye fields and prefrontal association cortex but also from both motor and premotor cortex. Even small HRP injections into the superficial layers of the superior colliculus yielded labelled cells in the agranular cortex (area 6) of the anterior bank of the arcuate sulcus. After large collicular injections affecting all layers, labelled cells were found in both motor and premotor cortex. This projection appeared to be topographically organized. Injections into the anterolateral parts of the superior colliculus labelled cells that were distributed within the presumed finger-hand--arm-shoulder representation, whereas after more caudal injections labelled cells occurred more in the presumed arm-trunk representation. The supplementary motor cortex was not found to contain labelled cells. The corticotectal cells in the motor cortex differed from those in the premotor cortex in their size distribution; the former being small, the latter both small and large. The functional significance of the motor and premotor input into the superior colliculus for sensory, and particularly visual, guidance of movements is discussed in view of a collicular role in the extrapersonal space representation and of its possible participation in steering arm and hand movements.     

Distribution of cells of origin of the corticotrigeminal projections to the nucleus caudalis of the spinal trigeminal complex in the cat. A horseradish peroxidase (HRP) study

The cortical distribution of cells of origin of the corticotrigeminal projections to the nucleus caudalis of the cat was examined using the method of retrograde axonal transport of horseradish peroxidase (HRP). After injections of HRP into the nucleus caudalis, labeled cells were distributed densely in the anterior suprasylvian gyrus, the coronal gyrus, and the ventral part of the anterior sigmoid gyrus, and moderately in the rostral part of the anterior ectosylvian gyrus on the contralateral side. In the anterior suprasylvian gyrus, the distribution extended rostrocaudally from the lateral ansate sulcal level to about 4.0 mm caudal to this level and mediolaterally throughout the convex of the anterior suprasylvian gyrus. All cortical labeled cells were pyramidal cells of various sizes in layer V.     

Brainstem projections to the phrenic nucleus: A HRP study in the cat

Brainstem neurones which project to the phrenic nucleus were identified using retrogradely transported horseradish peroxidase (HRP) as a marker. Following iontophoretic injection of HRP into the phrenic nucleus, labelled cells were encountered throughout large areas of the medulla and pons, but occurred with characteristic high densities in those regions known to contain phasic respiratory neurones: namely, the ventrolateral solitary tract nucleus (vl-NTS), known as the dorsal respiratory group (DRG), the ambiguus complex or ventral respiratory group (VRG) and the parabrachial pontine nuclei (BCM-KF). In 12 cats a total of 1540 cells was identified within these regions, the relative contralateral and ipsilateral contributions were respectively 72%:28% (vl-NTS), 65%:35% for the ambiguus complex, and 5%:95% (BCM-KF). In addition, labelled cells, predominantly ipsilateral, were observed in the pontine and medullary reticular formation and the vestibular nuclei. The labelled cells of the DRG had round, oval or triangular perikarya. Their mean soma diameter was 18.3 micrometers. The HRP-positive cells of the VRG had slightly larger somas (mean 21.2 micrometers) and they were fusiform and triangular. The neurones labelled in the BCM-KF nuclei were more heterogeneous with a mean soma size of 14.9 micrometers. The bilateral projections to the phrenic nucleus from the DRG and the VRG, and the predominantly ipsilateral projection from the BCM-KF are discussed in relation to current electrophysiological and autoradiographic findings.     

Callosal connections of the somatic sensory areas II and IV in the cat

The homotopic and heterotopic callosal connections in the forelimb representations of the second (SII) and fourth (SIV) somatic sensory areas of cats were investigated by means of the axonal transport of horseradish peroxidase (HRP) in conjunction with microelectrode recording. The tracer was injected in the electrophysiologically identified hand and/or digit zone of SII (six cats) or SIV (four cats). The homotopic area in the contralateral hemisphere was explored with microelectrodes in five animals (three injected in SII and two in SIV) to map neuronal receptive fields. The aim was to correlate in the same experimental case the topography of labelled callosal neurons with the physiological map of the forelimb. Labelled cells and recording sites were plotted on planar maps reconstructed with the aid of a computer from serial coronal sections from the anterior ectosylvian gyrus. After SII injections, labelled callosal neurons were observed throughout the forelimb representation in the contralateral area, but in the tangential plane their distribution was uneven. Each somatotopic zone composing the forelimb map, that is, the arm, hand, and digit zones, contained several subzones in which callosal neurons were either dense or rare. Microelectrode explorations showed that receptive fields mapped from callosal and relatively acallosal subzones representing the same body part were similar in extent and location. After SIV injections, labelled callosal neurons were observed throughout the forelimb and proximal body representation of the contralateral area. Although slight regional variations in the density of labelled cells were apparent, no subzones bare of callosal labelling were observed in SIV. In both SII and SIV, callosal neurons were concentrated mainly in layer III, but a significant number was also evident in the infragranular layers. After HRP injections in the digit zone of SII or SIV, labelled cell bodies were also observed in heterotopic areas of the contralateral hemisphere. Most of these neurons were clustered in the medial bank of the coronal sulcus and in two other heterotopic cortical regions lying, respectively, in the anterior suprasylvian sulcus and in the lateral branch of the ansate sulcus. Some callosal cells interconnecting SII and SIV were also labelled. The results show that the distal forelimb zones in SII and SIV are callosally connected with the respective homotopic zones and with several somatosensory fields located heterotopically in the contralateral hemisphere.     

Cells of origin of propriospinal connections to cat lumbosacral gray as determined with horseradish peroxidase

Propriospinal cells projecting to the lumbosacral spinal cord of cat were identified using the technique of retrograde transport of horseradish peroxidase (HRP). An injection technique was used to preferentially label cells via their terminals rather than through damaged axons of passage. Large volumes (1 to 5 μl) of 35% HRP were slowly ejected from a micropipet penetrating the cord dorsum midway between the dorsal root entry zone and the midline, while the pipet was slowly withdrawn from a depth of 1500 to 2000 μm below the cord surface. This technique resulted in diffusion of HRP throughout the gray matter on the side of the injection and, usually, sperad to the contralateral gray matter. HRP-labeled cells were observed from C1 to S3 after injections in segments L3-S3. Significant differences were seen in the rostrocaudal distributions of labeled cells after injections at different lumbosacral levels: specifically, L6-L7 injections resulted in less labeling of cells at cervical and high thoracic levels than injections that included segments L3-L5 or S1-S3.     

Nongeniculate afferents to striate cortex in macaques

Horseradish peroxidase (HRP) was injected in relatively massive amounts to cover most, or portions, of opercular striate cortex in four macaques. Absence of transcallosal or circumventricular labelling, plus discrete and consistent retrograde labelling in other areas in the four cases, assured the validity and specificity of the observations. Numerous labelled cells in regions directly bordering striate cortex, however, were excluded from the analysis because of the possibility of uptake consequent to physical diffusion. With this exception, all labelled cells were counted at roughly 2-mm intervals for one case with extensive unilateral injection of HRP. Even excluding the closely circumstriate population, the totals indicate that more than 30% of the afferent input to striate cortex arises from nongeniculate sources. Four areas of neocortex together make up about one-fourth of the total afferents: superior temporal sulcus 17.1%; inferior occipital area, 6.1%; intraparietal sulcus, 0.4%; and parahippocampal gyrus, 0.3%. Other areas projecting to striate cortex include claustrum, pulvinar, nucleus paracentralis, raphé system, locus coeruleus, and the nucleus basalis of Meynert. Cells of the latter were particularly striking with their very heavy uptake of HRP, and, even in cases of minimal effective injection, were scattered throughout an extensive area from the posterior edge of the globus pallidus passing rostrally beyond the chiasm and into the nucleus of the diagonal band. On the basis of their distribution and known cholinergic affinity, it is argued that this group also includes the cells labelled in and around lateral hypothalamus and cerebral peduncle, and that as a whole the group constitutes a cholinergic counterpart of the diffusely projecting monoaminergic systems. It seems possible that the basalis projection at first follows a fornical-subcallosal pathway to reach striate cortex via callosoperforant fibers.     

Cortical neurons projecting to the cervical and lumbar enlargements of the spinal cord in young and adult rhesus monkeys

Cortical neurons projecting to cervical and lumbar segments of the spinal cord in five young and one adult monkeys were identified using the retrograde transport method following multiple unilateral injections of horseradish peroxidase (HRP) into the anterior horn at cervical and lumbar levels of the spinal cord. Somatotopically organized labeled neurons were found in the precentral and postcentral gyri, the rostral half of both the medial and dorsal aspects of area 5, the cingulate sulcus within the medial aspect of area 6, and the second somatosensory area within the lateral sulcus. All HRP-positive neurons were confined to cortical layer V or to a depth corresponding to the fifth layer in regions where delineation of cortical layers was obscured due to freezing or sectioning artifacts. Although cross-sectional areas of labeled neurons varied widely within each field, more large labeled neurons were present in the leg than in the arm subdivision of the precentral gyrus. HRP-positive neurons in the first and second somatosensory areas as well as those in the medial aspect of area 5 were of medium size, and those in the primary and supplementary motor areas as well as those within the dorsal aspect of area 5 were of medium or larger size.     

Prefrontal granular cortex of the rhesus monkey. I. Intrahemispheric cortical afferents.

In 6 adolescent rhesus monkeys, unilateral injections of horseradish peroxidase (HRP) were made into 6 regions on the convexity of the prefrontal granular cortex.The afferents to each zone were considered with respect to whether they were local afferents (from adjacent frontal areas) or distal afferents (from outside frontal lobe). The strongest input onto prefrontal granular cortex comes from the temporal lobe and especially areas in and around the superior temporal gyrus. Area 10 in the frontal pole region receives input primarily from area 22 in the superior temporal gyrus and dorsal portion of the superior temporal sulcus. That portion of area 46 above the principal sulcus receives input primarily from area 22 in the upper bank of the superior temporal sulcus while area 46 below the principal sulcus has input from the insula of the superior temporal sulcus and area 21 in the lower bank of the superior temporal sulcus. The cortex within the concavity of the acurate sulcus differs in that the dorsal half (including areas 46 and 8a) receives input primarily from the dorsal bank and to a lesser degree the insula of the superior temporal sulcus while the ventral portion of this region including areas 45 and 46 receives input primarily from the lower bank of the superior temporal sulcus, inferior temporal gyrus and insula of the superior temporal sulcus. Input was noted from cingulate areas 23 and 24 to all 6 injected regions while retrosplenial cortex was noted to project to all but one of the injected regions, i.e. area 10. In addition, some labeled neurons were seen in area 7 after injections into area 46 and some were also seen in the inferior temporal gyrus and parahippocampal region after injections into the arcuate region. Finally, labeled neurons were noted in area 19 after injections into the ventral portion of the prefrontal granular cortex bounded by the arcuate sulcus.The HRP-positive neurons that comprised the intrahemispheric cortical afferents to prefrontal granular cortex were located primarily in layer iii. They were pyramidal in shape and ranged in size from small to medium. These neurons were found to be distributed in a horizontal band in which the number of labeled neurons waxed and waned, or they were distributed in a patchy or clumped manner. The possibility that both patterns of distribution represent a vertical or columnar organization to these afferent neurons is discussed.