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Differential spinal projections from the forelimb areas of the rostral and caudal subregions of primary motor cortex in the cat.

We used anterograde transport of WGA-HRP to examine the topography of corticospinal projections from the forelimb areas within the rostral and caudal motor cortex subregions in the cat. We compared the pattern of these projections with those from the somatic sensory cortex. The principal finding of this study was that the laminar distribution of projections to the contralateral gray matter from the two motor cortex subregions was different. The rostral motor cortex projected preferentially to laminae VI-VIII, whereas caudal motor cortex projected primarily to laminae IV-VI. Confirming earlier findings, somatic sensory cortex projected predominantly to laminae I-VI inclusive. We found that only rostral motor cortex projected to territories in the rostral cervical cord containing propriospinal neurons of cervical spinal segments C3-4 and, in the cervical enlargement, to portions presumed to contain Ia inhibitory interneurons. We generated contour maps of labeling probability on averaged segmental distributions of anterograde labeling for all analyzed sections using the same algorithm. For rostral motor cortex, heaviest label in the dorsal part of lamina VII in the contralateral cord was consistently located in separate medial and lateral zones. In contrast, no consistent differences in the mediolateral location of label was noted for caudal motor cortex. To summarize, laminae I-III received input only from the somatic sensory cortex, while laminae IV-V received input from both somatic sensory and caudal motor cortex. Lamina VI received input from all cortical fields examined. Laminae VII-IX received input selectively from the rostral motor cortex. For motor cortex, our findings suggest that projections from the two subregions comprise separate descending pathways that could play distinct functional roles in movement control and sensorimotor integration.

Animals↗

Supramedullary projections to the dorsal and ventral divisions of the paramedian reticular nucleus in the cat.

Injections of combined lectin-conjugated and unconjugated horseradish peroxidase were made in the dorsal (d) and ventral (v) divisions of the paramedian reticular nucleus (PRN), a precerebellar relay nucleus, of the cat. The origins of supramedullary afferent projections to the PRN were identified in the pons, midbrain and cerebral cortex using the transverse plane of section. The data indicate a segregation of input from a number of sites to the dPRN and vPRN. The interstitial nucleus of Cajal projects bilaterally to the dPRN and predominantly to the ipsilateral side. The vPRN receives only a unilateral projection from the ipsilateral nucleus of Cajal. Major afferent projections to the vPRN arise from the ipsilateral nucleus of Darkschewitsch and the intermediate layer of the contralateral superior colliculus. Neither of these sites projected to the dPRN. The raphe nuclei and medial reticular formation of the pons and midbrain contribute a moderate input to both divisions of the PRN. A moderate bilateral cerebral cortical projection arises from the first somatomotor area (SMI). The ventral coronal and anterior sigmoid gyri project mainly to the dPRN and vPRN respectively. Smaller afferent projections arise from the posterior sigmoid gyri and area 6 of Hassler and Mühs-Clement (1964) in the medial wall of the anterior sigmoid gyrus. Inputs from the accessory oculomotor nuclei, tectal regions and the first somatomotor cortex suggest a role in postural control for the PRN which may underlie its involvement in mediating orthostatic reflexes.

Animals↗

Specialized subregions in the cat motor cortex: anatomical demonstration of differential projections to rostral and caudal sectors.

Ipsilateral cortico-cortical and thalamo-cortical projections to the cat motor cortex were determined from the locations of retrogradely labeled neurons following single small intracortical injections of HRP in area 4 gamma. These projections were also examined by studying the distribution of anterogradely transported axonal label following multiple injections of HRP or of tritiated amino acids in areas 1-2 of SI and in area 2pri (SII). The number of retrogradely labeled cells in areas 1-2 and in area 2pri differed markedly between HRP injection sites located in the precruciate (anterior sigmoid gyrus) and postcruciate (posterior sigmoid gyrus) subregions of area 4 gamma. These associational projections from primary and secondary somatosensory cortices were dense to postcruciate subregions but weak to the precruciate subregions. The associational projections from areas 1-2 and from area 2pri to the postcruciate subregion of area 4 gamma were topographically organized, but no clear topographic organization could be demonstrated for the precruciate projection. Anterograde terminal labeling following injection of either HRP or tritiated amino acids into areas 1-2 and area 2pri confirmed the preferential projection of somatosensory cortex to the postcruciate subregion of motor cortex. The projection from somatosensory areas 1-2 was uniform over its terminal field, but that from area 2pri was more patchy and complex. HRP injections in area 4 gamma gave rise to lamellae of labeled neurons in the ventrolateral nucleus of thalamus (VL). A topographic relationship was found between the site of injection and the location of the lamella of labeled neurons. The percentage of retrogradely labeled neurons in the shell zone surrounding the border of the ventrolateral nucleus and the ventrobasal complex (VB) was greater following postcruciate than precruciate injections, whereas fewer retrogradely labeled neurons were found in central lateral nucleus (CL) after postcruciate injections than after precruciate injections. These observations support the hypothesis that differential cortical and thalamic projections to different subregions of area 4 gamma may give rise to the different physiological properties of neurons observed in these subregions (Vicario et al. 1983; Martin et al. 1981).

Afferent Pathways↗

The distribution of ipsilaterally and contralaterally projecting ganglion cells in the retina of the pigmented rabbit.

The retinal distribution of ipsilaterally and contralaterally projecting ganglion cells has been determined in the rabbit using both degeneration and horseradish peroxidase tracing techniques. Contralaterally projecting ganglion cells are present throughout the retinas, while ipsilaterally projecting ganglion cells are confined to a 3.0-3.5 mm wide strip adjacent to the temporal retinal margin. Thus, in this temporal strip both ipsilaterally and contralaterally projecting cells intermingle, while at more nasal locations all ganglion cells project contralaterally. Each of the contra- and ipsilaterally projecting populations comprises ganglion cells with soma diameters representing the full range present in the rabbit retina. However, a relatively large proportion of the ipsilaterally projecting ganglion cells have large somata (greater than or equal to 20 micrometer). Large ganglion cells are most numerous in the rabbit's temporal retina and have previously been described as reaching their peak density at the large cell node, just above the temporal end of the visual streak (Provis 1979). The large cell node lies immediately temporal to the nasal border of the strip of retina in which ipsilaterally projecting cells are located. It is possible that this specialization in the region of retina which observes the binocular visual field plays a particular role in binocular vision for the rabbit.

Afferent Pathways↗

Observations on the secondary vestibulocerebellar projections in the macaque monkey.

The distribution of retrogradely labeled cells in the nuclei of the vestibular nuclear complex following injections of horseradish peroxidase in various parts of the cerebellar cortex (except the nodulus and paraflocculus) has been mapped in the macacus rhesus monkey. In the main the findings correspond to those made in other mammalian species (cf. Table 1). The flocculus receives afferents bilaterally from the superior, medial and descending vestibular nucleus, group y, the interstitial nucleus of the vestibular nerve and also from the abducent nucleus. The projection to the posterior vermis (lobules VIII and IX), especially to lobule IX, is more abundant than that to lobules VI-VII. The projection to the anterior lobe vermis appears to be modest. Evidence for projections to the cerebellar hemispheres was not obtained. Whether the lateral vestibular nucleus projects to the cerebellum in the macaque is uncertain. The regular occurrence of weakly labeled cells among heavily labeled ones suggests that many of the cerebellar projecting cells may have axonal branches passing to other destinations. The findings lend support to the notion that there are precise topical relations within the entire secondary vestibulocerebellar projection. For example, in the medial nucleus the sites of origin of fibers to the flocculus and uvula are different. Surprisingly, many cells in group z were found to project to the uvula and - to a lesser extent - to lobule VIII. The group z may, therefore, not be a pure relay nucleus in a spinothalamic pathway, as generally assumed. The rather marked cerebellar projection of the abducent nucleus, especially to the flocculus, is of interest for the analysis of cerebellar control of eye movements in the macaque.

Abducens Nerve↗

Functional properties of neurons in the cat gracile nucleus that project to the dorsal accessory olive.

This study evaluated the functional properties of neurons in the gracile nucleus that project to the dorsal accessory portion of the inferior olive (DAO) and compared these with properties of other efferents from the dorsal column nuclei (DCN). Projection neurons were identified in anesthetized cats by microstimulation within DAO. They were further tested to insure that they could not be antidromically activated by stimulation in the medial lemniscus. Forty percent of the DAO projection neurons failed to respond to any form of natural stimulation. Of the 60% that did, most: 1) required stimuli of intensities greater than those needed to activate peripheral mechanoreceptors but less than those needed to excite peripheral nociceptors; 2) had receptive fields on the contralateral hindlimb or tail; and 3) could not follow peripheral stimuli presented at rates greater than 20 Hz. The conduction velocities of DAO projection neurons averaged only 4 m/s. In contrast, most of the neurons with axons in the medial lemniscus responded to very light cutaneous stimuli and followed peripheral stimuli presented at rates up to 100 Hz. Their average conduction velocity was 14 m/s. The low conduction velocity of the DAO projection neurons is unusual for DCN but is consistent with published descriptions of DAO. The limited responsiveness of the DAO projection neurons is, however, not uncharacteristic of DCN but places this projection at one end of the spectrum of DCN efferents, with the diencephalic projection at the other end. It is proposed that the DCN provide a filtered (and possibly movement-modulated) signal to the inferior olive for use in the control of movement.

Action Potentials↗

Convergence of cortico- and cuneopontine projections onto components of the pontocerebellar system in the rat: an anatomical and electrophysiological study.

Previous studies in the rat have demonstrated that corresponding peripheral tactile and somatosensory cortical inputs converge within the granule cell layer of various cerebellar lobules and further that descending corticopontine projections from the forelimb sensory cortex (FLSCx) partially overlap with the projection zones of ascending basilar pontine afferents from nucleus cuneatus (NC). The present study employed anatomical and electrophysiological procedures to determine whether cortical and dorsal column nuclear afferent projections converge on pontine neurons that, in turn, provide mossy fiber input to the granule cell layer of the paramedian lobule (PML), i.e., that portion of the rodent cerebellum shown to receive forelimb peripheral inputs. The combination of the orthograde and retrograde axonal transport of horseradish peroxidase (HRP) conjugated to wheat germ agglutinin (WGA) was used light microscopically to demonstrate that orthogradely labeled projections from injections of the FLSCx and NC converged with ponto-paramedian projection neurons that were retrogradely labeled from injections of the PML. These studies were also repeated in conjunction with ablations of either the FLSCx or NC which resulted in the ultrastructural identification of degenerating, as well as WGA-HRP labeled axonal boutons of these pontine afferent projections thus confirming that such projections actually formed synaptic contacts with the retrogradely labeled pontoparamedian projection neurons. Single unit recording analyses of neurons in the ventromedial region of the basilar pons following combined electrical stimulation of various regions of the sensorimotor cortex and the contralateral body surface indicated that approximately 40% of all cells recorded responded to electrical stimulation of corresponding regions of the cortex and periphery, particularly the FLSCx and the forepaw. Natural cutaneous stimuli applied to the forepaw that also elicited responses in these same groups of basilar pontine neurons and were associated with relatively small receptive fields. Taken together, these observations indicate that the previously observed convergence of peripheral and somatosensory cortical inputs within the granule cell layer of the cerebellar cortex may be at least partially organized at the level of the basilar pons.

Action Potentials↗

Collateralization of cerebellar efferent projections to the paraoculomotor region, superior colliculus, and medial pontine reticular formation in the rat: a fluorescent double-labeling study.

Collateralization of cerebellar efferent projections to the oculomotor region, superior colliculus (SC), and medial pontine reticular formation (mPRF) was studied in rats using fluorescent tracer substances. In one group, True Blue (TB) was injected into the oculomotor complex (OMC), including certain paraoculomotor nuclei and supraoculomotor ventral periaqueductal gray (PAG), and Diamidino Yellow (DY) was injected into the medial pontine reticular formation (mPRF) or pontine raphe. The largest number of single-TB-labeled (paraoculomotor-projecting) cells was observed in the medial cerebellar nucleus (MCN) and posterior interposed nucleus (PIN), whereas the largest number of single-DY-labeled (mPRF-projecting) cells was in the MCN. Double-TB/DY-labeled cells were present in the caudal two-thirds of the MCN, suggesting that some MCN neurons send divergent axon collaterals to the paraoculomotor region and mPRF. In another group, TB was injected into the SC and DY into the mPRF. The largest number of single-TB-labeled (SC-projecting) cells was in the PIN, although a considerable number of cells was observed in the caudal MCN, and ventral lateral cerebellar nucleus (LCN). Single-DY-labeled (mPRF-projecting) neurons were primarily located in the central and ventral MCN, but were also present in the lateral anterior interposed (AIN) and in the LCN. Double-TB/DY-labeled neurons were observed in the caudal two-thirds of the MCN and in the central portion of the LCN. The most significant new findings of the study concerned the MCN, which not only contained neurons that projected independently to the paraoculomotor region, SC, and mPRF, but also contained a considerable number of cells which collateralized to project to more than one of these nuclei. The possibility that the MCN projects to the supraoculomotor ventral PAG (containing an oculomotor interneuron system) and to the mPRF, which in the cat and monkey contain neural elements essential to the production of saccadic eye movements, is discussed. The anatomical findings suggest that the MCN in the rat plays an important role in eye movement.

Animals↗

A Golgi study on the neuronal organization of the interhemispheric cortex in the mouse. I. Projection neurons.

Projection neurons in the interhemispheric cortex (IHC) of the mouse were studied by the rapid Golgi method. Five layers were discerned in the IHC. Projection neurons in layer I had stellate or piriform cell bodies with dendrites which were distributed in layers I and II. The cell bodies of projection neurons in layer II were fusiform, piriform, triangular or stellate in shape. Fine axons of these neurons sent collaterals mainly to layer IV. Projection neurons in layer III were medium-sized pyramidal, and small spindle cells. Basal dendrites of the former neurons were distributed mainly in layer III, while those of the latter neurons extended into layer IV. Projection neurons in layer IV were largely pyramidal, medium-sized pyramidal, medium-sized fusiform, and small cells. In the large pyramidal cells, the basal dendrites were distributed mainly in layer IV, and the apical dendrites extended into layer I. The axons of these neurons sent collaterals to all cortical layers. In layer V, spindle and small stellate projection neurons were observed. All apical dendrites of projection neurons in layers I-III extended into layer I, whereas some apical dendrites of projection neurons in layers IV and V did not reach layer I.

Animals↗

The cost of developing site-specific environmental regulations: evidence from EPA's project XL.

The flagship of the Environmental Protection Agency's regulatory reinvention initiative, Project XL has been touted as a regulatory blueprint for a site-specific, performance-based pollution-control system, but widespread complaints about the costs of the program beg the question of whether the costs of tailoring regulations to individual facilities are manageable. To address this question, this paper presents original survey data on a sample of 11 XL projects. We find that the fixed costs of putting in place XL agreements are substantial, averaging over $450,000 per firm. While stakeholder negotiations are widely cited as the principal source for these costs, we find that they actually arise mainly from interaction between participating facilities and the EPA. Moreover, EPA management problems are perceived by our survey respondents as having inflated project development costs. Finally, we find that the key factors that explains differences in costs across XL projects are the scope and complexity of the project proposal. These findings suggest that Project XL favors large firms that can afford to pay significant project development costs, that EPA management problems must be resolved to reduce costs, and that there may be a significant economic bias against complex and innovative proposals--precisely the type of proposals that Project XL was designed to foster in order to improve the efficiency of the regulatory system.

Commerce↗

A population of pedal-buccal projection neurons associated with appetitive components of Aplysia feeding behavior.

Backfills of the cerebral-buccal connective (CBC) of Aplysia californica revealed a cluster of five to seven pedal-buccal projection neurons in the anterolateral quadrant of the ventral surface of each pedal ganglion. Intra- and extracellular recordings showed that the pedal-buccal projection neurons shared common electrophysiological properties and synaptic inputs. However, they exhibited considerable heterogeneity with respect to their projection patterns. All pedal-buccal projection neurons that were tested received a slow excitatory postsynaptic potential from the ipsi- and contralateral cerebral-pedal regulator (C-PR) neuron, a cell that is thought to play a key role in the generation of a food-induced arousal state. Tests were conducted to identify potential synaptic follower neurons of the pedal-buccal projection neurons in the cerebral and buccal ganglia, but none were detected. Finally, nerve recordings revealed projections from the pedal-buccal projection neurons in the nerves associated with the buccal ganglion. In tests designed to determine the functional properties of these peripheral projections, no evidence was obtained supporting a mechanosensory or proprioceptive role and no movements were observed when they were fired. It is proposed that peripheral elements utilized in consummatory phases of Aplysia feeding may be directly influenced by a neuronal pathway that is activated during the food-induced arousal state.

Animals↗

Efferent projections of the gracile nucleus in the cat.

The efferent projections of different portions of the gracile nucleus in the cat were studied using both autoradiographic and degeneration tracing methods. The results suggest that there are two aspects to the functional organization of these projections. First, the somatotopic organization of the gracile n. (GR) is maintained, but inverted, by the topographic organization of its projections to VPL1. Fibers from the lateral portions of GR terminate medially in VPL1; fibers from the dorsal portions terminate ventrally. These fibers, especially those from the middle and caudal portions of GR, terminate in dense, precisely located groups of clusters. Dorsally located clusters in VPL1 (predominantly from middle-ventral portions of GR) are significantly smaller than ventrally located clusters (predominantly from middle-dorsal portions). The second aspect of this organization, involving the projections both to VPL1 and to other brain stem targets, is that some kind of functionally relevant sorting process appears to occur as fibers leave different portions of the gracile n. The afferent projections of the rostral (GRr) and middle-ventral portions (GRmv) of the gracile n. are different from those from the other portions of the nucleus. Projections to VPL1 from GRr are less dense, less likely to form clusters, less clearly topographically organized, and extend further rostrally and dorsally in VPL1 than those from the rest of GR. The clusters are small, like those from GRmv. Similarly, although all portions of GR project to several other brain stem regions, these projections appear to be derived preferentially from GRr and/or GRmv. These brain stem regions involve certain portions of the inferior olive, inferior and superior colliculi, red n., zona incerta, pretectum, thalamic posterior group and the H field of Forel. This dual organization of efferent connectivity is similar to that of the cuneate n.20, and is consistent with many of the differences in cytoarchitecture, afferent connectivity and response properties of cells within different portions of the dorsal column nuclei.

Animals↗

Midbrain projections to the trigeminal, facial and hypoglossal nuclei in the opossum. A study using axonal transport techniques.

It has been proposed (see Berntson and Micco for review) that circuits intrinsic to the midbrain play an important role in the elaboration and control of behaviors involving the motor nuclei of the trigeminal, facial and hypoglossal nerves (e.g. defense, threat, attack); but because of technical problems, it has been difficult to analyze their organization. Using the horseradish peroxidase technique we have localized those midbrain neurons which project to each of the above nuclei and by using the autoradiographic method we have plotted the intranuclear distribution of their axons. Using both techniques, we have seen that mesencephalic projections to oral-facial motor nuclei strongly favor the nucleus of the facial nerve. Cells ventral to the cerebral aqueduct, including the ventral periaqueductal gray, the interstitial nucleus of Cajal, the nucleus of Darkshchewitsch and the rostral oculomotor nucleus provide major midbrain-facial projections in the opossum. Their axons terminate densely and bilaterally within areas innervating auricular muscles and to a lesser extent, the platysma sheet. The projection to the caudal auricular area of the facial complex is particularly dense. Neurons within and dorsal to the red nucleus project to regions of the contralateral facial nucleus reported to supply buccolabial, zygomatic and cervical musculature. There is also a minor tectal projection to the facial nucleus. Direct projections to the hypoglossal nuclei also arise within the periaqueductal gray and interstitial nucleus, but if such regions influence the motor trigeminal nucleus, it is mainly by way of dendrites that extend outside the nucleus or by at least one synaptic delay. The mesencephalic nucleus of the trigeminal nerve, however, projects strongly to the motor trigeminal nucleus. These data are discussed in light of their possible functional significance.

Animals↗

Topographic specificity of aberrant cerebellorubral projections following neonatal hemicerebellectomy in the rat.

Anterograde transport of horseradish peroxidase-wheat germ agglutinin (HRP-WGA) was used to examine the topographic specificity of ascending cerebellar efferent projections in adult rats which were hemicerebellectomized at birth. The results were compared to similar cerebellar projections in unlesioned adults. HRP-WGA placement in the nucleus interpositus of control rats resulted in a dense projection of labeled fibers which decussated in the midbrain, caudal to the red nucleus. In the red nucleus, dense terminal labeling was confined to the magnocellular region, while retrogradely labeled rubrocerebellar neurons were present throughout both parvo- and magnocellular areas. Similar HRP-WGA placements in the nucleus lateralis gave rise to fewer labeled fibers which terminated in the parvocellular red nucleus. In addition to the cerebellorubral projection, other areas of terminal labeling included the mid-brain reticular formation, nucleus parafascicularis prerubralis, zona incerta, fields of Forel and ventral thalamus. In neonatally lesioned adults, aberrant cerebellorubral and cerebellothalamic projections were observed deflecting ipsilaterally at the decussation of the normal contralateral projection. Topographic specificity of the aberrant ipsilateral cerebellorubral projection mirrored that of the normal contralateral fibers. In addition, an ipsilateral projection from the cerebellum could be followed rostral to the red nucleus, to terminate in the ipsilateral ventral thalamus. Lesioned animals also demonstrated marked cell loss in the red nucleus contralateral to the hemicerebellectomy.

Animals↗

The organization of neurons in the nucleus of the lateral lemniscus projecting to the superior and inferior colliculi in the rat.

The topographic organization of neurons in the dorsal nucleus of the lateral lemniscus (DNLL) which project to the superior and inferior colliculi was studied using the retrograde horseradish peroxidase (HRP) and the fluorescent double labeling methods. Neurons projecting to the superior colliculus (SC) are situated in the rostral portion of the DNLL, whereas those to the inferior colliculus (IC) are found in the caudal area of this nucleus. These two portions are completely separated from each other and no neurons projecting to both the SC and the IC are observed. In the dorsolateral part of the rostral portion of the DNLL, neurons projecting to the ipsilateral SC are found, whereas neurons projecting to the contralateral SC are located in the central to medial part of the nucleus, but no neurons sending collateral axons to both sides of the SC were observed. Neurons located in the central part of the caudal area of the DNLL project to the ipsilateral IC and neurons in the lateral and medial parts project contralaterally to the IC. Some of the neurons in the caudal part of the DNLL have divergent axonal branching projecting to both sides of the IC. In the ventral nucleus of the lateral lemniscus, labeled neurons were observed only when the HRP was injected into the ipsilateral IC.

Animals↗

Zonal organization of olivocerebellar projections to the uvula in rabbits.

Olivocerebellar projections to the uvula were studied by means of retrograde axonal transport of horseradish peroxidase (HRP) in pigmented rabbits. The distribution pattern of labeled cells in the inferior olive was compared among cases following large- and microinjections of HRP into the uvula. Findings indicate topographically organized projections to longitudinally oriented zones. There are at least 6 zones in the rabbit's uvula. The caudal part of the nucleus beta projects contralaterally to a most medially located zone (caudal beta zone). The rostral part of the nucleus beta projects to a little more laterally located zone (rostral beta zone) at a distance of about 1 mm from the midline of the uvula. The caudolateral part of the MAO projects to a zone (caudolateral MAO zone) located laterally to the rostral beta zone. The dorsomedial cell column projects to a zone (dorsomedial cell column zone) located in the intermediate part of the uvula at about 2 mm from the midline. The rostrolateral part of the MAO projects to the most lateral zone (rostrolateral MAO zone) of the uvula. Finally, the ventral lamella of the PO projects to a zone (ventral lamella of PO zone) located between the rostrolateral MAO zone and the dorsomedial cell column zone.

Animals↗

Cortical projection patterns of magnocellular basal nucleus subdivisions as revealed by anterogradely transported Phaseolus vulgaris leucoagglutinin.

The present paper deals with a detailed analysis of cortical projections from the magnocellular basal nucleus (MBN) and horizontal limb of the diagonal band of Broca (HDB) in the rat. The MBN and HDB were injected iontophoretically with the anterograde tracer Phaseolus vulgaris leucoagglutinin (PHA-L). After immunocytochemical visualization of labeled efferents, the distribution of projections over the cortical mantle, olfactory regions and amygdala were studied by light microscopy. Based on differences in cortical projection patterns, the MBN was subdivided in anterior, intermediate and posterior portions (MBNa, MBNi and MBNp). All subdivisions maintain neocortical projections and are subject to an anterior to posterior topographic arrangement. In the overall pattern, however, the frontal cortex is the chief target. Furthermore, all MBN parts project to various regions of meso- and allocortex, which are progressively more dense when the tracer injection is more anteriorly placed. The most conspicuous finding, however, was a ventrolateral to dorsomedial cortical projection pattern as the PHA-L injection site moved from posterior to anterior. Thus, the posterior MBN projects predominantly to lateral neo- and mesocortex while the anterior MBN sends more fibers to the medial cortical regions. Furthermore, the MBNa is a source of considerable afferent input to the olfactory nuclei and as such should be regarded as a transition to the HDB. The HDB, apart from projecting densely to olfactory bulb and related nuclei, maintains a substantial output to the medial prefrontal cortical regions and entorhinal cortex, as well. Comparison of young vs aged cases indicate that aging does not appear to have a profound influence on cortical innervation patterns, at least as studied with the PHA-L method.

Acetylcholinesterase↗

Organization of cutaneous primary afferent fibers projecting to the dorsal horn in the rat: WGA-HRP versus B-HRP.

Primary afferent projections from cutaneous afferents in the forelimb and hindlimb digits to the dorsal horn (DH) were examined using 4 tracers: (1) 25% free horseradish peroxidase (HRP), (2) 2.5% wheat-germ agglutinin conjugated to horseradish peroxidase (WGA-HRP), (3) a mixture of 25% free HRP and 2.5% WGA-HRP (WGA-HRP/HRP) or (4) 0.1% HRP conjugated to cholera toxin (B-HRP). The tracer was injected intracutaneously into the digits. Three to 4 days later, the rats were perfused transcardially, transverse sections (60-microns thick) were cut and the HRP was reacted using the tetramethyl benzidine (TMB) method. The location of the label was reconstructed by camera lucida drawings. In rats which received an injection of HRP alone, no label was detected in the DH. Rats injected with WGA-HRP had projection patterns similar to those injected with WGA-HRP/HRP. Patterns of labelling with WGA-HRP differed markedly from those with B-HRP. WGA-HRP labelled cutaneous afferents projecting to Rexed's laminae I-III, with the densest label in lamina II; in contrast, B-HRP labelled cutaneous afferents projecting to laminae II-V, with the densest label in laminae III-IV. These results indicate that, for cutaneous primary afferents projecting to the DH, WGA-HRP and B-HRP labelled different subpopulations of fibers, with the B-HRP-labelled subpopulation biased toward afferents of larger diameter. Rostrocaudally, the extent of the densest fiber projections, whether labelled by WGA-HRP or by B-HRP, was essentially the same, but the extent of the less densely labelled projections was much greater with B-HRP than with WGA-HRP. Comparisons of the projection maps from each of the five digits, using either WGA-HRP or B-HRP, indicated that, as seen in transverse sections through the DH, there was extensive overlapping among the labelled cutaneous afferent fibers from adjacent, or even non-adjacent digits.

Afferent Pathways↗