Orientation topography of layer 4 lateral networks revealed by optical imaging in cat visual cortex (area 18)

The functional specificity of corticocortical connections with respect to the topography of orientation selectivity was studied by optical imaging of intrinsic signals and bulk injections of fluorescent latex beads (green and red) and biocytin into layer 4. The distributions of retrogradely labelled cells and anterogradely labelled axon terminals were histologically reconstructed from all cortical laminae, and the resulting anatomical maps compared with the optically imaged functional maps. Layer 4 injections produced extensive horizontal labelling up to 2-3 mm from the injection centres albeit without the clear patchy pattern described after layer 2/3 injections (Gilbert & Wiesel 1989, J. Neurosci., 9, 2432-2442; Kisvárday et al. 1997, Cerebral Cortex, 7, 605-618). The functional (orientation) distribution of the labelled projections was analysed according to laminar location and lateral spread. With regard to the former, no major difference in the orientation topography between supragranular- (upper tier), granular- (middle tier) and infragranular (lower tier) layers was seen. Laterally, proximal and distal projections were distinguished and further dissected into three orientation categories, iso- (+/- 30 degrees ), oblique- (+/- 30-60 degrees ) and cross-orientations (+/- 60-90 degrees ) with respect to the orientation preference at the injection sites. The majority of distal connections (retrograde and anterograde) was equally distributed across orientations (35.4% iso-, 33.7% oblique-, and 30.9% cross-orientations) that are equivalent with a preponderance to dissimilar orientations (oblique- and cross-orientations, 64.6%). In one case, distal excitatory and inhibitory connections could be morphologically distinguished. For both categories, a marked bias to dissimilar orientations was found (excitatory, 63.7%; inhibitory, 86.6%). Taken together, these results suggest that the long-range layer 4 circuitry has a different functional role from that of the iso-orientation biased (52.9%, Kisvárday et al. 1997, Cerebral Cortex, 7, 605-618) layer 2/3 circuitry, and is perhaps involved in feature difference-based mechanisms, e.g. figure ground segregation.     

Transplant connectivity in the rat cerebral cortex. A carbocyanine study

Previous reports suggest that the specificity of connectivity between cortical areas and their related thalamic nuclei is established during development. We have used heterotopic E15, E17 and E19 transplants to determine at which embryonic age the cortex is mature enough to establish such specific connections. Carbocyanine DiI or Fluoro-Gold were used for labelling transplants. Efferent cortico-cortical, callosal and collicular connections were observed when DiI was applied in fixed tissue. Thalamic nuclei were labelled when DiI was used 'in vivo' or the transplants were labelled with Fluoro-Gold. The connections of the transplants were those corresponding to the area in which the transplants were placed. This suggests that up to E19 the embryonic cortex is too immature to define the specific thalamo-cortical connections.     

Corticocortical connections within the primary somatosensory cortex of the rat

Corticocortical connections within the primary somatosensory (SI) cortex of rat were investigated by using discrete injections of retro- and orthogradely transported neuroanatomical tracers (including HRP, WGA, PHA-L, and 3H-leucine). Tangential and vertical connections were defined with respect to the cytoarchitectonic divisions within the rat SI, specifically: (1) the "granular zones" (GZs), characterized by their dense layer IV granular aggregates, which receive the majority of direct ventroposterior (VP) thalamocortical terminations, (2) the "perigranular zones" (PGZs), the less-granular cortical matrix just surrounding the GZs, and (3) the "dysgranular zones" (DZs), the larger dysgranular regions lying centrally within and just lateral to the SI. Receptive fields recorded in the granular zones are small and discrete, whereas in the perigranular zones and especially in dysgranular zones they exhibit complex sensory convergence. A major aim of this study was to determine whether the pattern of intracortical connectivity within the SI is compatible with these observed physiological differences. In general, the perigranular and dysgranular zones contained more profuse systems of corticocortical connections than did the granular zones. For example, discrete tracer injections in the perigranular zones produced "walls" of labelling throughout the adjacent perigranular zones, while adjacent granular zones were relatively empty. Nevertheless, the granular zones were filled with dendritic branches of neurons in adjacent perigranular zones. Since these dendrites could presumably receive direct VP thalamocortical contacts, they represent one path through which this thalamic sensory information might be transmitted to the perigranular zones. Further transmission to the dysgranular zones might be subserved by a topographically organized system of reciprocal interconnections that was found between the perigranular zones and dysgranular zones. In coronal sections, labelling produced by relatively distant injections of either retro or orthograde tracers generally appeared in a columnar distribution, and was localized in perigranular zones and dysgranular zones. Within these zones, orthograde labelling consisted of vertically oriented axons emitting collateral sprays of terminals in all layers. Retrograde neuronal labelling (composed almost exclusively of pyramidal cells) was greatest in supragranular layers. Proximal to the injection site, labelling tended to spread out from these columns into supra- and infragranular layers in adjacent granular zones.(ABSTRACT TRUNCATED AT 400 WORDS)     

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.     

Topography of the afferent connectivity of area 17 in the macaque monkey: a double-labelling study

The various structures afferent to area 17 (or V1) of the macaque monkey have widely differing retinotopic organizations. It is likely that these differences are reflected in the topographic organizations of the projections from these structures to area V1. We have investigated this issue by placing side-by-side injections of two retrograde fluorescent tracers, fast blue and diamidino yellow, in V1. By examining the extent of mixing of the two populations of singly labelled cells and the presence of doubly labelled cells, in different structures, we have characterized the topography of each projection in terms of the size of its axonal arborization and the amount of convergence and divergence. The afferents from the lateral geniculate nucleus (LGN) and from the pulvinar are organized in a point-to-point fashion. The maximum extent of axonal arborization of these afferents is 0.5 mm and these projections demonstrate little scatter (i.e., neighboring LGN neurons project to adjacent regions of V1). The other two subcortical structures examined, the claustrum and the intralaminar nuclei, demonstrate a much larger scatter and wider axonal arborizations in their projections to V1 than do the LGN and pulvinar. Two-dimensional reconstructions were made of the distribution of labelled neurons in extrastriate cortical areas. Using the separation between patches of labelled cells and transitions in myelin-stained sections, we have identified seven separate cortical regions containing labelled cells. Two of these can be identified as area V2 and the middle temporal visual area (MT). Three other regions correspond to areas V3, V3A and V4t. Finally, two more regions of labelling have been distinguished that belong to area V4. These results demonstrate that, at least within the central 6 degrees of visual field, all the presently known extrastriate visual cortical areas project to V1. This result is interesting in view of the fact that only a few extrastriate cortical areas are reported to receive afferents from V1. Three groups of cortical areas can be distinguished on the basis of the characteristics of their cortical connections to V1. The first group contains area V2, V3, and the posterior region of V4. These areas project to V1 with infra- as well as supragranular layer neurons and show limited axonal arborization and scatter in the projection. The second group consists of two regions of labelling in the superior temporal sulcus corresponding to V4t and MT and another on the annectant gyrus (V3A). These regions contain almost exclusively infragranular labelling and show wide axonal arborization and scatter in their projections to V1.(ABSTRACT TRUNCATED AT 400 WORDS)     

Relationship Between Lateral Inhibitory Connections and the Topography of the Orientation Map in Cat Visual Cortex

The functional and structural topography of lateral inhibitory connections was investigated in visual cortical area 18 using a combination of optical imaging and anatomical tracing techniques in the same tissue. Orientation maps were obtained by recording intrinsic signals in regions of 8.4–19 mm². To reveal the inhibitory connections provided by large basket cells, biocytin was iontophoretically injected at identified orientation sites guided by the pattern of surface blood vessels. The axonal and dendritic fields of two retrogradely labelled large basket cells were reconstructed in layer III. Their axonal fields extended up to 1360 μm from the parent somata. In addition to single basket cells, the population of labelled basket cell axons was also studied. For this analysis anterogradely labelled basket axons running horizontally over 460–1280 μm from the core of an injection site in layer III were taken into account. The distribution of large basket cell terminals according to orientation preferences of their target regions was quantitatively assessed. Using the same spatial resolution as the orientation map, a frequency distribution of basket cell terminals dependent on orientation specificity could be derived. For individual basket cells, the results showed that, on average, 43% of the terminals provided input to sites showing similar orientation preferences (±30°) to those of the parent somata. About 35% of the terminals were directed to sites representing oblique-orientation [±(30–60)°], and 22% of them terminated at cross-orientation sites [±(60–90)°]. Furthermore, the possible impact of large basket cells on target cells at different distances and orientation preferences was estimated by comparing the occurrence of orientation preferences with the occurrence of basket terminals on the distance scale. It was found that a basket cell could elicit iso-orientation inhibition with a high impact between 100–400 and 800–1200 μm strong cross-orientation inhibition at ～400–800 μm, and oblique-orientation inhibition between 300–500 and 700–900 μm from the parent soma. The non-isotropic topography of large basket axons suggests a complex function for this cell class, possibly including inhibition related to orientation and direction selectivity depending on the location of the target cells and possible target selectivity.     

Anatomical organization of retinotectal afferents in the cat: an autoradiographic study.

The distribution of retinotectal afferents has been studied by autoradiography in 4 adult cats. The findings suggest that crossed and uncrossed retinal fibers terminate in a striking cluster-and-sheet pattern that varies systematically with respect to the retinotopic map of the colliculus. Following unilateral eye injection, labelling was most pronounced in the contralateral colliculus but a suprising volume of label appeared on the ipsilateral side in all cases in the form of dense clumps of silver grains separated by sparsely labelled zones. The contralateral projection appeared densest in the most superficial of the 3 laminae of the stratum griseum superficiale; appreciable labelling was present also in the middle lamina at all survival times used (23-72 h). Near the area centralis representation labelling in both contralateral tiers weakened markedly and local gaps appeared densest in the more dorsal band. Elsewhere, labelling in this dorsal band was generally dense, though sharply interrupted at the optic disc representation and in a curious, elongated lateral zone at mid-collicular levels. In the caudal half of the binocular zone rarefications or 'holes', about 200 mum wide, appeared in the more ventral tier between more densely labelled zones of roughly similar width. On the ipsilateral side, labelling was sparse or absent at the rostral and caudal collicular poles, and was also weak in the region of the area centralis representation save for occasional very superficial grain-clusters. Farther caudally, however, prominent approx. 200 mum wide 'puffs' of label marked the middle lamina of the superficial gray layer. The puffs were most regular in shape in the caudal half of the ipsilateral zone and these were spaced at roughly 200 mum intervals. Puffs lateral to the horizontal meridian representation tended to lie more dorsal than those medial to this line and some of the most lateral puffs at mid-collicular levels invaded the upper lamina of the superficial gray layer. The optic disc representation was marked by a column of label extending through the upper and middle laminae. Similar experiments in cat fetuses suggest that these staggered--and possible even complementary--patterns of crossed and uncrossed retinotectal projection are innate: ipsilateral 'puffs' of labelling and contralateral 'holes' appear in the superior colliculus at least one week before term, as does the ipsilateral filling-in and contralateral gap at the optic disc representation. These observations suggest that in the cat, a vertical as well as horizontal organization may characterize the superficial layers of the superior colliculus. The additional finding of a similar, interrupted puff-like pattern of labelling in the stratum griseum medium following injections in the region of the substantia nigra makes it likely that a somewhat comparable cluster-and-sheet organization may exist also in the deep collicular layers.     

Connections of the primary auditory cortex in the common marmoset, Callithrix jacchus jacchus

The afferent and efferent connections of the primary auditory cortex (AI) of common marmosets were traced following small injections of wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) made at best frequency (BF)-defined sites in the AI. After the injections the animals remained anesthetized for 15-23 hours; they were then perfused transcardially with fixative and the brains were processed for WGA-HRP reaction product. Examination of the disposition of labelled material revealed the following results. First, patches of terminal labelling, and to a lesser extent retrograde labelling, were found outside the injection site in the ipsilateral cortex rostral and caudal to the AI. Second, the region of the contralateral cortex corresponding to the injection site contained labelled terminals throughout the depth of the cortex; labelled neurons were found in the middle layers. Third, in each experiment a discrete region of the medial geniculate body (MG) contained retrogradely labelled neurons interspersed with anterogradely labelled terminals. These regions had a banded appearance, were found in the dorsal and rostral half of the MG, and shifted in location progressively dorsalward as the injection site BF increased. The presence of projection zones rostral and caudal to the AI of marmosets, and the disposition of the MG sources of projection in relation to BF, are similar to observations made on other New World monkeys. The ipsilateral corticocortical projections confirm electrophysiological evidence suggesting the existence of auditory fields rostral and caudal to the AI. The thalamocortical auditory system of the marmoset appears relatively simple, with a comparatively undifferentiated MG projecting to a cortical auditory system dominated by a large AI.     

Topography of retinal representation in the rabbit cortex: An experimental study using transneuronal and retrograde tracing techniques

The cortical representation of the retina in rabbits was studied autoradiographically using transneuronal transport of tritiated proline and tritiated fucose. Injection of one eye produced cortical labelling on both sides. There was no indication of ocular dominance columns. On the contralateral side, the labelling was found in cortical layers II–IV, and partly also in layer VI of area striata and the caudal part of area peristriata, and in layer IV of most of area occipitalis. In addition, there was a patch of labelling in layers II–IV of the temporal cortex. On the ipsilateral side, the labelling was restricted to layer IV in a strip of cortex comprising area striata and area occipitalis along their common border. The cortical labelling in area striata and occipitalis was interpreted as evidence for a direct geniculo-cortical projection. This was confirmed by demonstration of retrogrately labelled nerve cells in the lateral geniculate nucleus produced by small injections of horseradish peroxidase into area occipitalis and area striata. In experiments with intraocular injections of tritiated adenosine, transneuronal cortical labelling was much weaker and more diffuse. In addition to the neuropil, nerve cell bodies and glial cells were also labelled.     

Organization in the somatosensory sector of the cat's thalamic reticular nucleus

This study describes the organization of cells in the thalamic reticular nucleus (TRN) that project to the somatosensory part of the dorsal thalamus in the cat. Injections of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) and fluorescent dyes were made into the ventrobasal complex (VB) and the medial division of the posterior complex (POm) of the thalamus. The resultant retrograde labelling in TRN was analyzed. Large injections of a tracer in VB label many reticular cells that are restricted to a centroventral, or somatosensory, sector of TRN. Small injections of a tracer in VB produce narrow zones of labelled cells in this sector. In reconstructions these zones resemble thin “slabs,” which lie parallel to the plane of TRN along its oblique rostrocaudal dimension and occupy only a fraction of its thickness. In comparisons of the zones of labelled cells in TRN resulting from tracer injections in different nuclei of VB, inner cells, intermediate cells, and outer cells across the thickness of TRN project to the ventral posteromedial, the medial division of the ventral posterolateral, and the lateral division of the ventral posterolateral nuclei, respectively. Furthermore, shifts in injected areas along the dorsoventral dimension of VB produce similar shifts in zones of labelled cells in TRN. Thus, reticular cells form an accurate map on the basis of their connections with VB. Large injections of a tracer in the ventral subdivision of POm label many reticular cells that are also restricted to the centroventral sector of TRN. Small injections of a tracer in ventral POm produce broad zones of labelled cells in this sector. In comparisons of the zones of labelled cells in TRN resulting from tracer injections in different regions of ventral POm, cells that project to these regions are scattered across the thickness of TRN and occupy overlapping territories. Large injections of a tracer in either VB or ventral POm also label cells in a restricted centroventral region of the perireticular nucleus. Double injections of different tracers in VB and ventral POm produce many cells in TRN that are labelled from both of these dorsal thalamic structures and fewer cells that are labelled from only one or the other of these structures. These results indicate that there is a dual organization in the projections of cells in the somatosensory sector of TRN to dorsal thalamus: Projections to VB are topographically organized whereas those to ventral POm lack a topographical organization. Furthermore, both of these mapped and nonmapped projections can arise from single reticular cells in the somatosensory sector. © 1996 Wiley-Liss, Inc.     

Projections from the Cerebral Cortex to the Red Nucleus of the Guinea-pig. A Retrograde Tracing Study

The origin and the topographic distribution of corticorubral (CR) projections in the guinea-pig were studied by using the retrograde axonal transport of a tracer, colloidal gold-labelled, enzymatically inactive horseradish peroxidase conjugated to wheat-germ agglutinin (WGAapoHRP - Au), which was injected in the red nucleus (RN). It was found that the bulk of the CR projections arise from layer V neurons of the agranular frontal cortex in both its medial (Agm) and lateral (Agl) subdivisions; in the Agm labelled neurons are preferentially located in the upper part of layer V, whereas in the Agl they are more concentrated in the central band of the layer. Fewer projections originate from areas of the granular parietal and the agranular cingulate and retrobulbar cortices. CR projections have a bilateral origin, with a large ipsilateral predominance. The pattern of retrograde cortical labelling observed after injection of WGAapoHRP - Au in different portions of the RN indicates that CR projections are distributed throughout the entire rostrocaudal extent of the nucleus, but are slightly more concentrated in the rostral parvocellular area. The morphological arrangement of CR projections in the guinea-pig, as demonstrated in the present study, shows several analogies with other mammals. The functional characteristics of the cortical areas in which CR neurons are located indicate that CR projections may play a significant role in the central organization of movement.     

Connections between the reticular nucleus of the thalamus and pulvinar-lateralis posterior complex: A WGA-HRP study

The present study utilises the capacity of wheat germ agglutinin-conjugated horseradish peroxidase to label both afferent and efferent projections from selected regions of the thalamic reticular nucleus (TRN) to the pulvinar lateralis-posterior complex (Pul-LP) of the cat. Fourteen injections into the TRN located between anterior-posterior levels 8.5 and 4.5 were analysed. The projection of the TEN to the Pul-LP complex is roughly organised in a topographic manner and is not widespread within the thalamus. Anterograde labelling in the Pul-LP extended rostrocaudally with a slight oblique dorsoventral orientation. Projections to the medial LP were predominantly but not exclusively from rostral areas of TRN, while projections to the lateral LP were largely from caudal areas of the TRN. Projections to other areas of the Pul-LP were sparse. The connections between TRN and Pul-LP were reciprocal, although the distribution of labelled cells and anterograde labelling was not completely overlapping. Reciprocal connections with the dorsal lateral geniculate nucleus were largely with the C-laminae and the medial interlaminar nucleus. The results are discussed with reference to the corticothalamic projections and the visuotopy of the Pul-LP. © 1995 Wiley-Liss, Inc.     

Afferent and efferent connections of the parabigeminal nucleus in cat revealed by retrograde axonal transport of horseradish peroxidase

Afferent and efferent connections of the parabigeminal nucleus (PBG) of the cat have been demonstrated by means of horseradish peroxidase (HRP) tracing technique. Following HRP injection in the PBG, labelled cells were observed mainly in the deep layers of the ipsilateral superior colliculus (SC). The other labelled structures were the prepositus hypoglossi complex (PH), the ventral nucleus of the lateral geniculate body (LGV), the locus coeruleus, the cuneiform nucleus, the periaqueductal gray and the dorsomedial hypothalamic area. Efferent projections of the PBG were investigated by HRP injection in SC, LGV, PH, hypothalamus and in some acoustic relays, i.e. medial geniculate body and inferior colliculus. Only the PBG-SC projection appeared to be well systematized. The positive labelling of the PBG following injection of LGV and hypothalamus is discussed in terms of the specificity of the injection. The absence of afferent and efferent connections of the PGB with any acoustic relay tends to exclude this nucleus from the auditory system in contrast to previous suggestions. On the basis of the close reciprocal PBG-SC connections a possible role of the PBG within visuomotor tectal function is proposed.     

Fibre organization of the monkey's optic tract: I. Segregation of functionally distinct optic axons

The fibre organization of the monkey's optic tract was examined by implanting pellets of horseradish peroxidase into different locations within the tract, or into the superior colliculus and pretectum. Retinae were examined for the distribution, size, and morphological types of retrogradely labelled ganglion cells; optic tracts were examined for the distribution of anterogradely and retrogradely labelled axonal profiles; and lateral geniculate nuclei were examined for the distribution of anterogradely labelled processes within distinct geniculate laminae. Localized implants in the optic tract produced retrograde labelling of ganglion cells across wide regions of the retinal surface. The maximum density of labelled cells was always substantially less than the total ganglion cell density known to be present at those retinal loci. Distinct retinal ganglion cell types were labelled from differing regions within the optic tract: implants into the deep (dorsal) portion of the tract, far removed from the outer, pial, surface, retrogradely labelled predominantly P beta retinal ganglion cells, whereas implants into the superficial (ventral), subpial, part of the tract retrogradely labelled primarily the other retinal ganglion cell types, i.e., the P alpha, P gamma, and P epsilon cells. Within any given class of axon, there is a mapping of the centroperipheral retinal axis across the deep-to-superficial dimension of the tract, but this retinotopy is extremely coarse. Anterograde labelling of axonal terminations within the lateral geniculate nucleus showed a corresponding specificity for distinct geniculate laminae, the deep implants labelling the parvocellular laminae, superficial implants labelling the magnocellular laminae. Implants into the visual centres of the midbrain produced retrograde axonal labelling rostral to the lateral geniculate nucleus only in the superficial part of the optic tract. These results demonstrate that the monkey's optic tract is not a simple topographic mapping of retinal eccentricity. Rather, the primary organizational principle is that of a segregation of functionally distinct optic axon classes. As fibre order in the mammalian optic tract is also a chronological index of axonal arrival during development, the present results provide specific predictions about the temporal order of ganglion call genesis and axonal addition within the visual pathway. They also provide an anatomical basis for the functionally selective visual impairments that may arise following local damage to the optic tract in humans.     

Columnar specificity of intrinsic horizontal and corticocortical connections in cat visual cortex

A prominent and stereotypical feature of cortical circuitry in the striate cortex is a plexus of long-range horizontal connections, running for 6-8 mm parallel to the cortical surface, which has a clustered distribution. This is seen for both intrinsic cortical connections within a particular cortical area and the convergent and divergent connections running between area 17 and other cortical areas. To determine if these connections are related to the columnar functional architecture of cortex, we combined labeling of the horizontal connections by retrograde transport of rhodamine-filled latex microspheres (beads) and labeling of the orientation columns by 2-deoxyglucose autoradiography. We first mapped the distribution of orientation columns in a small region of area 17 or 18, then made a small injection of beads into the center of an orientation column of defined specificity, and after allowing for retrograde transport, labeled vertical orientation columns with the 2-deoxyglucose technique. The retrogradely labeled cells were confined to regions of orientation specificity similar to that of the injection site, indicating that the horizontal connections run between columns of similar orientation specificity. This relationship was demonstrated for both the intrinsic horizontal and corticocortical connections. The extent of the horizontal connections, which allows single cells to integrate information over larger parts of the visual field than that covered by their receptive fields, and the functional specificity of the connections, suggests possible roles for these connections in visual processing.     

The inferior olivary connections to the cerebellum in the rat studied by retrograde axonal transport of horseradish peroxidase

Olivocerebellar projections were investigated in the rat using retrograde axonal transport of horseradish peroxidase. Discrete cell groups of the inferior olive were labelled, subsequent to injections in the paravermal region, the vermis, or the caudolateral hemisphere. Injections in the midrostrocaudal third of the paravermal area resulted in labelling of cells in the medial accessory olive (MAO), in cell group “b” at caudal levels, and in its lateral portion at mid-rostrocaudal levels. The rostral pole of the principal olive (PO), the dorsal accessory olive (DAO), and the dorsomedial cell column, were heavily labelled. By comparison, caudal paravermal injections resulted in labelling in the medial part of the mid-rostrocaudal levels of the MAO, but not in its caudal portion. The PO lamellae were labelled in their lateral half, excluding the lateral bend connecting them. Injections slightly lateral within this paravermal area gave no caudal MAO labelling, but did label cells in segments of both PO lamellae, medial to those in the previous case. From vermal injections, cell groups “b” and “c” of the caudal MAO were labelled, but no labelled cells were present in the PO. Subsequent to injections in the paramedian lobule, cells in the dorsal lamella of the PO were labelled. No cells of the MAO were labelled. These results are discussed in terms of specific labelling patterns and the general concepts of organization presently held for the Olivocerebellar system.     

The Topographic Organization and Axis of Projection within the Visual Sector of the Rabbit's Thalamic Reticular Nucleus

The organization of the visual field representation within the thalamic reticular nucleus (TRN) of the rabbit was studied. Focal injections of horseradish peroxidase (HRP) and/or [3H]proline were made into visuocortical areas V1 and V2 and the dorsal lateral geniculate nucleus (dLGN). The resultant labelling in the thalamus was analysed. A single injection in V1 or V2 results in a single zone of terminal label within the TRN that is restricted to the dorsocaudal part of the sheet-like nucleus. In comparisons of the zones of label following injections at two different cortical sites in V1, a medial to lateral shift in label across the thickness of the TRN sheet is accompanied by a medial to lateral shift in label in the dLGN; a dorsal to ventral shift in label within the plane of the TRN sheet is accompanied by a dorsal to ventral shift in label in the dLGN. Thus, like the dLGN the TRN receives a precise topographic projection from V1. In reconstructions from horizontal sections the zones of label within the TRN resemble 'slabs', which lie within the plane of the nucleus parallel to its borders. Thus, the slabs of visuocortical terminals and reticular dendrites are similarly oriented. As revealed by the orientation of the slabs, the lines of projection representing points in visual space are represented by the oblique rostrocaudal dimension of the TRN. Injections restricted to V1 produce terminal labelling that is confined to the outer two-thirds of the TRN across its thickness, whilst those involving V2 result in terminal labelling within the inner one-third of the nucleus. Thus, the adjacent cortical areas V1 and V2 project in a continuous fashion across the mediolateral dimension of the TRN. The organization of the map within the TRN, which was revealed by visuocortical injections, was confirmed by the pattern of retrograde labelling within the nucleus following geniculate injections of HRP. On the basis of these findings and those in other mammalian species, two major conclusions can be reached that alter our view of the TRN. First, rather than mapping onto the whole nucleus in a continuous fashion, the cortical projection to the TRN has significant discontinuities. Second, rather than integrating efferents from widespread cortical areas, the reticular dendrites are related to focal areas of cortex.     

Topography of developing thalamic and cortical pathways in the visual system of the cat

Adult patterns of connectivity could emerge during development by a process of selective elimination from an earlier, more widespread, connectivity. We have addressed this issue by examining the topography of developing projections to area 17 in the cat. At different postnatal ages, paired injections of the retrograde tracers diamidino yellow and fast blue were made in area 17. Interinjection separations were carefully controlled and the spatial distribution of the two populations of labelled neurones investigated. Projections to the striate cortex from the lateral geniculate nucleus, area 18, as well as connections intrinsic to area 17 were analysed quantitatively with a graphic method that uses a two-dimensional model of the projection. This allows two parameters of the projection to be calculated: the divergence (the spatial extent of area 17 contacted by an infinitely small region of an afferent structure) and the convergence (the extent of an afferent structure that projects to an infinitely small region of area 17). During postnatal development, the bulk of the connections making up the geniculostriate and corticocortical pathways showed no variation either in their convergence and divergence. However, the projection of area 18 to area 17 and the intrinsic area 17 connections (but not the geniculostriate projection) in the 3–15-day-old kittens were each found to contain a small subpopulation of widely scattered neurones with widespread axonal trajectories. These results, showing that many initially formed connections display a high degree of topographical order, are discussed in terms of the control mechanisms specifying axonal trajectories during development. © 1994 Wiley-Liss, Inc.     

Organization of projections from olfactory epithelium to olfactory bulb in the frog, Rana pipiens

One hypothesis for the coding of olfactory quality is that regions of the olfactory epithelium are differentially sensitive to particular odor qualities and that this regional sensitivity is conveyed to the olfactory bulb in a topographic manner by the olfactory nerve. A corollary to this hypothesis is that there is a sufficiently orderly connection between the epithelium and the olfactory bulb to convey this topographical coding. Thus we examined topography in the projection from epithelium to bulb in the frog, which has been the subject of numerous electrophysiological studies but has not yet been examined using modern neuroanatomical techniques. The tracer WGA-HRP was applied to the ventral or to the dorsal olfactory epithelium, or both. Anterograde transport of label to the olfactory bulb was seen after as few as 2 days; label was still present in the bulb as long as 21 days postinjection. In cases where WGA-HRP was applied to the entire epithelium, there was dense anterograde labelling of the ipsilateral olfactory bulb. In addition, a small medial portion of the contralateral bulb was labelled. Injections limited to either the ventral or dorsal epithelium produced patterns of anterograde labelling in the glomerular layer of the olfactory bulb, which varied with the size and location of the injection. With very large injections in either the dorsal or ventral epithelium, label appeared to be evenly distributed in the glomerular layer. With smaller injections in the ventral epithelium, there was heavier labelling in the lateral than in the medial portions of the glomerular layer, although light labelling was found in all regions of the glomerular layer. In contrast, injection sites restricted to the dorsal epithelium produced more anterograde labelling in the medial than lateral portions of the glomerular layer. These patterns extended throughout the dorsal-ventral extent of the bulb. Within the limits of the anterograde tracing technique used, we were unable to detect any systematic relationship between the pattern of labelling in the glomerular layer and the medial-lateral or rostral-caudal location of the injection site in either the ventral or dorsal epithelium. We conclude that in the frog, as in other amphibia, there is only a limited degree of topographic order between the epithelium and the olfactory bulb.