Auditory cortex projections target the peripheral field representation of primary visual cortex

The purpose of the present study was to identify projections from auditory to visual cortex and their organization. Retrograde tracers were used to identify the sources of auditory cortical projections to primary visual cortex (areas 17 and 18) in adult cats. Two groups of animals were studied. In the first group, large deposits were centered on the lower visual field representation of the vertical meridian located along the area 17 and 18 border. Following tissue processing, characteristic patterns of cell body labeling were identified in extrastriate visual cortex and the visual thalamus (LGN, MIN, & LPl). In auditory cortex, of the four tonotopically-organized regions, neuronal labeling was identified in the supragranular layers of the posterior auditory field (PAF). Little to no labeling was evident in the primary auditory cortex, the anterior auditory field, the ventral posterior auditory field or in the remaining six non-tonotopically organized regions of auditory cortex. In the second group, small deposits were made into the central or peripheral visual field representations of primary visual cortex. Labeled cells were identified in PAF following deposits into regions of primary visual cortex representing peripheral, but not central, visual field representations. Furthermore, a coarse topography was identified in PAF, with neurons projecting to the upper field representation being located in the gyral portion of PAF and neurons projecting to the lower field representation located in the sulcal portion of PAF. Therefore, direct projections can be identified from tonotopically organized auditory cortex to the earliest stages of visual cortical processing.     

Corticostriatal transformations in the primate somatosensory system. Projections from physiologically mapped body-part representations.

1. The basal ganglia of primates receive somatosensory input carried largely by corticostriatal fibers. To determine whether map-transformations occur in this corticostriatal system, we investigated how electrophysiologically defined regions of the primary somatosensory cortex (SI) project to the striatum in the squirrel monkey (Saimiri sciureus). Receptive fields in the hand, mouth, and foot representations of cortical areas 3a, 3b, and 1 were mapped by multiunit recording; and small volumes of distinguishable anterograde tracers were injected into different body-part representations in single SI areas. 2. Analysis of labeled projections established that at least four types of systematic remapping occur in the primate corticostriatal system. 1) An area of cortex representing a single body part sends fibers that diverge to innervate multiple regions in the putamen, forming branching, patchy fields that are densest in the lateral putamen. The fields do not form elongated cylindrical forms; rather, they are nearly as extended mediolaterally as they are rostrocaudally. 2) Cortical regions representing hand, mouth, and foot send globally somatotopic, nonoverlapping projections to the putamen, but regions with closely related representations (such as those of the thumb and 5th finger in area 3b) send convergent, overlapping corticostriatal projections. The overlap is fairly precise in the caudal putamen, but in the rostral putamen the densest zones of the projections do not overlap. 3) Regions representing homologous body parts in different SI cortical areas send projections that converge in the putamen. This was true of paired projections from areas 3a and 3b, and from areas 3b and 1. Thus corticostriatal inputs representing distinct somatosensory submodalities can project to the same local regions within the striatum. Convergence is not always complete, however: in the rostral putamen of two cases comparing projections from areas 3a and 1, the densest zones of the projections did not overlap. 4) All projections from SI avoid striosomes and innervate discrete zones within the matrix. 3. These experiments demonstrate that the somatosensory representations of the body are reorganized as they are projected from SI to the somatosensory sector of the primate putamen. This remapping suggests that the striatal representation of the body may be functionally distinct from that of each area of SI. The patchy projections may provide a basis for redistribution of somatosensory information to discrete output systems in the basal ganglia. Transformations in the corticostriatal system could thus be designed for modulating different movement-related programs.     

Topography of corticothalamic projections from the auditory cortex of the rat

Corticothalamic projections from cortical auditory field to the medial geniculate body (MG) in the rat were systematically examined by making small injections of biocytin in cortical area Te1. All injections, confined to 400 microm in diameter, resulted in two projections terminating in the ventral (MGV) and dorsal divisions (MGD) of the MG. The projections to the MGV were evidently topographic. The rostral and caudal portions of area Te1 projected to the ventromedial and dorsolateral parts of the MGV, respectively, forming narrow bands of terminal axons that extended in the mediolateral direction in the coronal plane of the MGV. The minimum dorsoventral width of the bands ranged approximately from 100 to 300 microm. Besides, the more rostral portion of area Te1 tended to project to the more rostral side of the MGV. The projections to the MGD consistently arborized in its ventral margin made up of the deep dorsal nucleus of the MGD. A similar weak topography along the rostrocaudal direction was observed in the projections to the MGD. Large terminals were occasionally found in the MGD after the injections involving cortical layer V. The distribution of large terminals also appeared topographic along with small terminals that were the major component of labeling. Collaterals of labeled axons produced slabs of terminal field in the thalamic reticular nucleus, which also exhibited a weak topography of distribution. These results provide insights into the structural basis of corticofugal modulations related to the tonotopic organizations in the cortex and MG.     

Efferent connections of "posterodorsal" auditory area in the rat cortex: implications for auditory spatial processing

We examined efferent connections of the cortical auditory field that receives thalamic afferents specifically from the suprageniculate nucleus (SG) and the dorsal division (MGD) of the medial geniculate body (MG) in the rat [Neuroscience 117 (2003) 1003]. The examined cortical region was adjacent to the caudodorsal border (4.8-7.0 mm posterior to bregma) of the primary auditory area (area Te1) and exhibited relatively late auditory response and high best frequency, compared with the caudal end of area Te1. On the basis of the location and auditory response property, the cortical region is considered identical to "posterodorsal" auditory area (PD). Injections of biocytin in PD revealed characteristic projections, which terminated in cortical areas and subcortical structures that play pivotal roles in directed attention and space processing. The most noticeable cortical terminal field appeared as dense plexuses of axons in area Oc2M, the posterior parietal cortex. Small terminal fields were scattered in area frontal cortex, area 2 that comprises the frontal eye field. The subcortical terminal fields were observed in the pontine nucleus, the nucleus of the brachium inferior colliculus, and the intermediate and deep layers of the superior colliculus. Corticostriatal projections targeted two discrete regions of the caudate putamen: the top of the middle part and the caudal end. It is noteworthy that the inferior colliculus and amygdala virtually received no projection. Corticothalamic projections terminated in the MGD, the SG, the ventral zone of the ventral division of the MG, the ventral margin of the lateral posterior nucleus (LP), and the caudodorsal part of the posterior thalamic nuclear group (Po). Large terminals were found in the MGD, SG, LP and Po besides small terminals, the major component of labeling. The results suggest that PD is an auditory area that plays an important role in spatial processing linked to directed attention and motor function. The results extend to the rat findings from nonhuman primates suggesting the existence of a posterodorsal processing stream for auditory spatial perception.     

Cytoarchitecture of the medial geniculate body and thalamic projections to the auditory cortex in the rufous horseshoe bat (Rhinolophus rouxi). I. Temporal fields

The auditory cortex in echolocating bats is one of the best studied in mammals, yet the projections of the thalamus to the different auditory cortical fields have not been systematically analyzed in any bat species. The data of the present study were collected as part of a combined investigation of physiological properties, neuroarchitecture, and chemoarchitecture as well as connectivity of cortical fields in Rhinolophus in order to establish a neuroanatomically and functionally coherent view of the auditory cortex in the horseshoe bat. This paper first describes the neuroanatomic parcellation of the medial geniculate body and then concentrates on the afferent thalamic connections with auditory cortical fields of the temporal region. Deposits of horseradish peroxidase and wheatgerm-agglutinated horseradish peroxidase were made into neurophysiologically characterized locations of temporal auditory cortical fields; i.e., the tonotopically organized primary auditory cortex, a ventral field, and a temporal subdivision of a posterior dorsal field. A clear topographic relationship between thalamic subdivisions and specific cortical areas is demonstrated. The primary auditory cortex receives topographically organized input from the central ventral medial geniculate body. The projection patterns to the temporal subdivision of the posterior dorsal field suggest that it is a "core" field, similar to the posterior fields in the cat. Projections to the ventral field arise primarily from border regions of the ventral medial geniculate body. On the whole, the organization of the medial geniculate body projections to the temporal auditory cortex is quite similar to that described in other mammals, including cat and monkey.     

Cytoarchitecture of the medial geniculate body and thalamic projections to the auditory cortex in the rufous horseshoe bat (<Emphasis Type="Italic">Rhinolophus rouxi)</Emphasis>

The auditory cortex in echolocating bats is one of the best studied in mammals, yet the projections of the thalamus to the different auditory cortical fields have not been systematically analyzed in any bat species. The data of the present study were collected as part of a combined investigation of physiological properties, neuroarchitecture, and chemoarchitecture as well as connectivity of cortical fields in Rhinolophus in order to establish a neuroanatomically and functionally coherent view of the auditory cortex in the horseshoe bat. This paper first describes the neuroanatomic parcellation of the medial geniculate body and then concentrates on the afferent thalamic connections with auditory cortical fields of the temporal region. Deposits of horseradish peroxidase and wheatgerm-agglutinated horseradish peroxidase were made into neurophysiologically characterized locations of temporal auditory cortical fields; i.e., the tonotopically organized primary auditory cortex, a ventral field, and a temporal subdivision of a posterior dorsal field. A clear topographic relationship between thalamic subdivisions and specific cortical areas is demonstrated. The primary auditory cortex receives topographically organized input from the central ventral medial geniculate body. The projection patterns to the temporal subdivision of the posterior dorsal field suggest that it is a “core” field, similar to the posterior fields in the cat. Projections to the ventral field arise primarily from border regions of the ventral medial geniculate body. On the whole, the organization of the medial geniculate body projections to the temporal auditory cortex is quite similar to that described in other mammals, including cat and monkey.     

Cortical convergence from different frequency domains in the cat primary auditory cortex

Primary auditory cortex (AI) has a tonotopic map consisting of orderly isofrequency (IF) bands, and cortical connections are commonly supposed to link domains preferring similar characteristic frequencies (CFs) within AI and in auditory association cortex. The interaction of different frequency channels, however, has not fully been understood in terms of anatomical substrates. Here, by injecting two anterograde tracers in different frequency domains of cat AI, without overlap of the injection cores, we attempted to relate the anatomical mapping of cortical outputs to physiologically defined fields in the auditory cortex. Consistent with previous studies, patches of labeled axon terminals were oriented largely along the IF axis. In regions distant from the injection sites, however, terminal patches were divergent in distribution. This divergence resulted in a complex geometry of partial overlap of projections originating from the two injection sites. The relative extent of the overlap tended to vary depending on the distance between the two injection sites. Physiological mapping for tonotopy across auditory fields revealed that projectional overlap was characteristic of dorsal AI and the dorsoposterior field and, to a lesser extent, in the secondary auditory field. Considering the differences in frequency representation in different AI IF bands, the anatomical convergence of projections tuned to different CFs could contribute to the spectral integration of sound components. Furthermore, the different extent of convergence in the functionally distinct fields might reflect field-specific processing of acoustic signals.     

Topographic organization of somatosensory corticotectal influences in cat

Using electrophysiological techniques, the present study demonstrated that substantial direct somatosensory cortical influences on the superior colliculus (SC) originate from three areas: a) SIV, b) para-SIV (the cortex adjacent to SIV but deeper in the anterior ectosylvian sulcus (AES) and for which no topography has yet been described), and c) the rostral suprasylvian sulcus. Influences also appeared to originate from SI and SII, but these may have been indirect. Detailed examination of the AES revealed that these corticotectal projections are topographically organized, and stimulation of a given cortical locus was observed to affect only those cells in the SC whose receptive fields overlapped those of cells at the stimulation site. A similar receptive-field register was found between the suprasylvian sulcus and the SC. Within this topographic pattern, considerable convergence was evident and an individual SC cell could be influenced from a surprisingly large cortical area. This was particularly evident within the representation of the forelimb. Thus, an SC cell with a receptive field covering the forelimb and paw could receive convergent input from many cortical cells with receptive fields covering all or restricted portions of this body region. Considerable corticotectal divergence also was observed within this general topographic scheme. For example, a given corticotectal site representing the digits sent projections to many different SC cells that included the digits within their receptive fields. These data are more consistent with a block-to-block than a point-to-point corticotectal projection. Somatosensory corticotectal projections excited only those SC cells that could also be activated by peripheral somatosensory stimuli. Similarly, the caudal AES, which contains auditory cells, excited only those SC cells activated also by peripheral auditory stimuli. Yet convergent influences from both auditory and somatosensory regions of the AES were observed in the SC cells that could be activated by both auditory and somatosensory stimuli. These data indicate that the AES is a major source of excitatory input to cells of the deep laminae of the SC. Since it is these deep laminae cells that project to premotor regions of the brain stem and the spinal cord, it is reasonable to suppose that the AES has a significant impact on the output signals of the SC that initiate the orientation responses to peripheral sensory stimulation.     

Topographical and laminar distribution of cortical input to the monkey entorhinal cortex

Hippocampal formation plays a prominent role in episodic memory formation and consolidation. It is likely that episodic memory representations are constructed from cortical information that is mostly funnelled through the entorhinal cortex to the hippocampus. The entorhinal cortex returns processed information to the neocortex. Retrograde tracing studies have shown that neocortical afferents to the entorhinal cortex originate almost exclusively in polymodal association cortical areas. However, the use of retrograde studies does not address the question of the laminar and topographical distribution of cortical projections within the entorhinal cortex. We examined material from 60 Macaca fascicularis monkeys in which cortical deposits of either (3)H-amino acids or biotinylated dextran-amine as anterograde tracers were made into different cortical areas (the frontal, cingulate, temporal and parietal cortices). The various cortical inputs to the entorhinal cortex present a heterogeneous topographical distribution. Some projections terminate throughout the entorhinal cortex (afferents from medial area 13 and posterior parahippocampal cortex), while others have more limited termination, with emphasis either rostrally (lateral orbitofrontal cortex, agranular insular cortex, anterior cingulate cortex, perirhinal cortex, unimodal visual association cortex), intermediate (upper bank of the superior temporal sulcus, unimodal auditory association cortex) or caudally (parietal and retrosplenial cortices). Many of these inputs overlap, particularly within the rostrolateral portion of the entorhinal cortex. Some projections were directed mainly to superficial layers (I-III) while others were heavier to deep layers (V-VI) although areas of dense projections typically spanned all layers. A primary report will provide a detailed analysis of the regional and laminar organization of these projections. Here we provide a general overview of these projections in relation to the known neuroanatomy of the entorhinal cortex.     

Microstructural organizational patterns in the human corticostriatal system

The axons that project into the striatum are known to segregate according to macroscopic cortical systems; however, the within-region organization of these fibers has yet to be described in humans. We used in vivo fiber tractography, in neurologically healthy adults, to map white matter bundles that originate in different neocortical areas, navigate complex fiber crossings, and project into the striatum. As expected, these fibers were generally segregated according to cortical origin. Within a subset of pathways, a patched pattern of inputs was observed, consistent with previous ex vivo histological studies. In projections from the prefrontal cortex, we detected a topography in which fibers from rostral prefrontal areas projected mostly to rostral parts of the striatum and vice versa for inputs originating in caudal cortical areas. Importantly, within this prefrontal system there was also an asymmetry in the subset of divergent projections, with more fibers projecting in a posterior direction than anterior. This asymmetry of information projecting into the basal ganglia was predicted by previous network-level computational models. A rostral-caudal topography was also present at the local level in otherwise somatotopically organized fibers projecting from the motor cortex. This provides clear evidence that the longitudinal organization of input fields, observed at the macroscopic level across cortical systems, is also found at the microstructural scale at which information is segregated as it enters the human basal ganglia.     

Topographical organization of the projections from physiologically identified areas of the motor cortex to the striatum in the rat.

The present study was undertaken to determine in the rat the topography of the neostriatal projections originating from the motor cortex. For that purpose, anterograde tracers (Phaseolus vulgaris leucoagglutinin: PHA-L; wheat germ agglutinin conjugated to horseradish peroxidase: WGA-HRP) were deposited in discrete cortical sites physiologically identified by microstimulation. Five major motor areas were considered in this study: the rostral (RFL) and caudal (CFL) forelimb areas, the hindlimb (HL) area, the vibrissae motor-frontal eye field (V-FEF) region and the jaw, lips and tongue (JLT) area (according to the nomenclature of Neafsey et al.). The results indicate that functionally different regions of the motor cortex project to different sectors of the caudate putamen (CPU). All 3 distinct limb areas RFL, CFL and HL project to the dorsolateral quarter of the CPU, V-FEF area projects to the dorsomedial quarter, whereas the JLT area projects to the ventrolateral quarter. The pattern of terminal labeling is relatively consistent, whatever the cortical area in which the tracer is deposited. This pattern is characterized by the presence of two or more labeled bands which are obliquely oriented along a ventrolateral-dorsomedial axis. Control experiments were also undertaken in which a retrograde tracer (WGA-HRP) was deposited in various neostriatal loci. The results are congruent with the findings of the anterograde study and further indicate that a given neostriatal sector receives projections from cytoarchitectonically different but functionally related regions of the neocortex. The somatotopic features of both motor and somatosensory corticostriatal projections appear to be in register. In addition, the striatal distribution of motor cortical fibers was compared in 6 experimental cases to the compartmental subdivision of the striatum in patches and matrix, following immunohistochemical localization of calbindin 28 kDa. The calbindin-immunoreactivity is extremely weak in the dorsolateral sector but is higher in the central and ventrolateral parts of the CPU. In these deep striatal regions receiving fibers from V-FEF, JLT and, to a lesser extent, from the limb areas, the cortical fibers are mostly directed to the matrix. The band-like organization of the projection from the motor cortex is correlated to the patch-matrix organization. The patches correspond to the bands of low density of terminal fibers and the matrix to the bands of high terminal density. The present results provide an anatomical basis to both electrophysiological and behavioral observations suggesting that functional distinctions can be established between subregions of the striatum.     

Neocortical connections in fetal cats

The major extrinsic projections to and from visual and auditory areas of cerebral cortex were examined in fetal cats between 46 and 60 days of gestation (E46-E60) using axonal transport of horseradish peroxidase either alone or in combination with tritiated proline. Projections to visual cortex from the dorsal lateral geniculate nucleus and lateral-posterior/pulvinar complex exist by E46, and those from the contralateral hemisphere, claustrum, putamen, and central lateral nucleus of the thalamus are present by E54-E56. In addition, cells in the medial geniculate nucleus project to auditory cortex by E55. At E54-E56 efferent cortical projections reach the contralateral hemisphere, claustrum, putamen, lateral-posterior/pulvinar complex and reticular nucleus of the thalamus. Cells in visual cortex also project to the dorsal and ventral lateral geniculate nuclei, pretectum, superior colliculus and pontine nuclei, and cells in auditory cortex project to the medial geniculate nucleus. Except for interhemispheric projections, all pathways demonstrated are ipsilateral, and projections linking cerebral cortex with claustrum, dorsal lateral geniculate nucleus and lateral-posterior/pulvinar complex are reciprocal. The reciprocal projections formed with the dorsal lateral geniculate nucleus, lateral-posterior/pulvinar complex and the claustrum show a greater degree of topological organization compared to the projections formed with the contralateral hemisphere and superior colliculus, which show little or no topological order. Therefore, the results of the present study show that the major extrinsic projections of the cat's visual and auditory cortical areas with subcortical structures are present by the eighth week of gestation, and that the origins and terminations of many of these projections are arranged topologically.     

Spatial organization of the corticopalladial projection system of dog brain

Spatial organization of corticopallidal projectional system was studied in 11 outbred dogs by method based on horse radish peroxidase transport. It was demonstrated that globus pallidum receives projections predominantly from neocortical zones (motor, premotor, somatosensory, parietal and auditory and from insular field of mesocortex. Mesocortical (prelimbic, orbital and insular) and allocortical (entorhinal, piriform and periamygdalar) including archicortex (subicular part of hippocampal formations) fields project onto ventral pallidum. Entopeduncular nucleus receives projections from neocortical zones (motor, premotor, somatosensory, parietal and auditory), mesocortex (prelimbic, orbital, insular and cingular fields) and allocortex (entorhinal and periamygdalar fields). The data obtained indicate specificity of distribution of cortical afferent projectional fibres in each of nuclei studied which allows to consider globes pallidum as motor zone and ventral pallidum as limbic zone of paladial complex. As projections from functionally different cortical fields were revealed in entopeduncular nucleus it may be suggested that this is the exact site for interaction of functionally different information, including the one received from the cortex.     

Convergence of unimodal and polymodal sensory input to the entorhinal cortex in the fascicularis monkey

The hippocampal formation is a key structure in memory formation and consolidation. The hippocampus receives information from different cortical and subcortical sources. Cortical information is mostly funneled to the hippocampus through the entorhinal cortex (EC) in a bi-directional way that ultimately ends in the cortex. Retrograde tracing studies in the nonhuman primate indicate that more than two-thirds of the cortical afferents to the EC come from polymodal sensory association areas. Although some evidence for the projection from visual unimodal cortex to the EC exists, inputs from other visual and auditory unimodal association areas, and the possibility of their convergence with polymodal input in the EC remains largely undisclosed. We studied 10 Macaca fascicularis monkeys in which cortical deposits of the anterograde tracer biotinylated dextran-amine were made into different portions of visual and auditory unimodal association cortices in the temporal lobe, and in polymodal association cortex at the upper bank of the superior temporal sulcus. Visual and auditory unimodal as well as polymodal cortical areas projected to the EC. Both visual unimodal and polymodal association cortices presented dense projections, while those from unimodal auditory association cortex were more patchy and less dense. In all instances, the projection distributed in both the superficial and deep layers of the EC. However, while polymodal cortex projected to all layers (including layer I), visual unimodal cortex did not project to layer I, and auditory unimodal cortex projected less densely, scattered through all layers. Topographically, convergence from the three cortical areas studied can be observed in the lateral rostral and lateral caudal subfields. The present study suggests that unimodal and polymodal association cortical inputs converge in the lateral EC, thereby providing the possibility for the integration of complex stimuli for internal representations in declarative memory elaboration.     

Processing of temporal information and the basal ganglia: new evidence from fMRI

Temporal information processing is a fundamental brain function, which might include central timekeeping mechanisms independent of sensory modality. Psychopharmacological and patient studies suggest a crucial role of the basal ganglia in time estimation. In this study, functional magnetic resonance imaging (fMRI) was applied in 15 healthy right-handed male subjects performing an auditory time estimation task (duration discrimination of tone pairs in the range of 1,000–1,400 ms) and frequency discriminations (tone pairs differing in pitch, around 1,000 Hz) as an active control task. Task difficulty was constantly modulated by an adaptive algorithm (weighted up-down method) reacting on individual performance. Time estimation (vs rest condition) elicited a distinct pattern of cerebral activity, including the right medial and both left and right dorsolateral prefrontal cortices (DLPFC), thalamus, basal ganglia (caudate nucleus and putamen), left anterior cingulate cortex, and superior temporal auditory areas. Most activations showed lateralisation to the right hemisphere and were similar in the frequency discrimination task. Comparing time and frequency tasks, we isolated activation in the right putamen restricted to time estimation only. This result supports the notion of central processing of temporal information associated with basal ganglia activity. Temporal information processing in the brain might thus be a distributed process of interaction between modality-dependent sensory cortical function, the putamen (with a timing-specific function), and additional prefrontal cortical systems related to attention and memory. Further investigations are needed to delineate the differential contributions of the striatum and other areas to timing. Electronic Publication     

Interconnections of auditory areas in the guinea pig neocortex

By studying the efferent projections of five auditory areas in the guinea pig cortex, we sought evidence that the larger fields can be divided into subareas based on unique patterns of cortical connections. Small extracellular injections of biocytin were made in combination with evoked potential mapping or single-unit analysis and histochemical determination of cortical landmarks. The two core fields, primary (AI) and dorsocaudal (DC), are partially surrounded by six adjacent belt areas, leaving two gaps: one at the rostral edge of AI and the other at the dorsal edge. All of the areas studied projected to their nearest neighbors, but AI was the only area to project to all seven of the other auditory areas. The caudal, high-frequency (more than 4 kHz) end of AI had different projections from the rostral, low-frequency (less than 1.5 kHz) end, and there was no evidence of connections between the two ends. Each end had separate dorsal and ventral projections. The two ends of AI may be working independently. By contrast, area DC had strong connections between its high- and low-frequency ends and it may be involved in auditory/visual integration. The dorsorostral belt (DRB) was subdivided into two zones on the basis of its projections: the more rostral part appears to overlap the second somatosensory area and be bimodal, while the caudal part has stronger auditory connections. The small belt area (area S) had separate physiological and anatomical properties from the rest of the rostral belt.     

Somatotopically arranged inputs from putamen and subthalamic nucleus to primary motor cortex

Employing retrograde transsynaptic transport of rabies virus, we investigated the organization of basal ganglia inputs to hindlimb, proximal and distal forelimb, and orofacial representations of the macaque primary motor cortex (MI). Four days after rabies injections into these MI regions, neuronal labeling occurred in the striatum and the subthalamic nucleus (STN) through the cortico-basal ganglia loop circuits. In the striatum, two distinct sets of the labeling were observed: one in the dorsal putamen, and the other in the ventral striatum (ventromedial putamen and nucleus accumbens). The dorsal striatal labeling was somatotopically arranged and its distribution pattern was in good accordance with that of the corticostriatal inputs, such that the hindlimb, orofacial, or forelimb area was located in the dorsal, ventral, or intermediate zone of the putamen, respectively. The distribution pattern of the ventral striatal labeling was essentially the same in all cases. In the STN, the somatotopic arrangement of labeled neurons was in register with that of corticosubthalamic inputs. The present results suggest that the cortico-basal ganglia motor circuits involving the dorsal putamen and the STN may constitute separate closed loops based on the somatotopy, while the ventral striatum provides common multisynaptic projections to all body-part representations in the MI.     

The auditory pathway in cat corpus callosum

The cortical auditory fields of the two hemispheres are interconnected via the corpus callosum. We have investigated the topographical arrangement of auditory callosal axons in the cat. Following circumscribed biocytin injections in the primary (AI), secondary (AII), anterior (AAF) and posterior (PAF) auditory fields, labelled axons have been found in the posterior two-thirds of the corpus callosum. Callosal axons labelled by small individual cortical injections did not form a tight bundle at the callosal midsagittal plane but spread over as much as one-third of the corpus callosum. Axons originating from different auditory fields were roughly topographically ordered, reflecting to some extent the rostro-caudal position of the field of origin. Axons from AAF crossed on average more rostrally than axons from AI; the latter crossed more rostrally than axons from PAF and AIL Callosal axons originating in a discrete part of the cortex travelled first in a relatively tight bundle to the telo-diencephalic junction and then dispersed progressively. In conclusion, the cat corpus callosum does not contain a sector reserved for auditory axons, nor a strictly topographically ordered auditory pathway. This observation is of relevance to neuropsychological and neuropathological observations in man.     

Restricted cortical termination fields of the midline and intralaminar thalamic nuclei in the rat.

The projections from the midline and intralaminar thalamic nuclei to the cerebral cortex were studied in the rat by means of anterograde tracing with Phaseolus vulgaris-leucoagglutinin. The midline and intralaminar nuclear complex taken as a whole projects to widespread, predominantly frontal, cortical areas. Each of the constituent thalamic nuclei has a restricted cortical projection field that overlaps only slightly with the projection fields of adjacent midline and intralaminar nuclei. The projections of the intralaminar nuclei cover a larger cortical area than those of the midline nuclei. The laminar distributions of fibres from individual midline and intralaminar thalamic nuclei are different and include both deep and superficial cortical layers. The parataenial, paraventricular and intermediodorsal midline nuclei each project to circumscribed parts of the prefrontal cortex and the hippocampal and parahippocampal regions. In the prefrontal cortex, the projections are restricted to the medial orbital, infralimbic, ventral prelimbic and agranular insular fields, and the rostral part of the ventral anterior cingular cortex. In contrast to the other midline nuclei, the rhomboid nucleus projects to widespread cortical areas. The rostral intralaminar nuclei innervate dorsal parts of the prefrontal cortex, i.e. the dorsal parts of the prelimbic, anterior cingular and dorsal agranular insular cortical fields, the lateral and ventrolateral orbital areas, and the caudal part of the ventral anterior cingular cortex. Additional projections are aimed at the agranular fields of the motor cortex and the caudal part of the parietal cortex. The lateral part of the parafascicular nucleus sends fibres predominantly to the lateral agranular field of the motor cortex and the rostral part of the parietal cortex. The medial part of the parafascicular nucleus projects rather sparsely to the dorsal part of the prelimbic cortex, the anterior cingular cortex and the medial agranular field of the motor cortex. Individual midline and intralaminar thalamic nuclei are thus in a position to directly influence circumscribed areas of the cerebral cortex. In combination with previously reported data on the organization of the midline and intralaminar thalamostriatal projections and the prefrontal corticostriatal projections the present results suggest a high degree of differentiation in the convergence of thalamic and cortical afferent fibres in the striatum. Each of the recently described parallel basal ganglia-thalamocortical circuits can thus be expanded to include projections at both the cortical and striatal levels from a specific part of the midline and intralaminar nuclear complex. The distinctive laminar distributions of the fibres originating from the different nuclei emphasize the specificity of the midline and intralaminar thalamocortical projections.