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

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

Nucleus sagulum: projections of a lateral tegmental area to the inferior colliculus in the cat

The nucleus sagulum, an area of the midbrain tegmentum, has been considered a component of a lateral tegmental system within the ascending auditory pathway to the thalamus. In this study, connections of the nucleus sagulum within the midbrain were investigated in adult cats. Tracing methods using anterograde and retrograde axonal transport of markers were employed. The nucleus sagulum was identified as a region of principally small neurons (261 +/- 79 micron2) at the margin of the midbrain and neighboring the nuclei of the lateral lemniscus. Injections of tritiated leucine in the nucleus sagulum labeled axons that ended in dense patches within the superficial layers of the caudal portion of the dorsal cortex of the inferior colliculus on the ipsilateral side. Retrograde experiments confirmed this connection. Other axonal projections labeled in the anterograde studies included fibers ending in the dorsomedial nucleus, the superficial layers of the dorsal cortex, and the rostral nucleus of the inferior colliculus with some bilateral distribution. Outside of the inferior colliculus, sagulum injections labeled other axons ending in the ventral intercollicular tegmentum on both sides and in a dorsal and rostral region of the contralateral nucleus sagulum that appeared contiguous with the dorsal nucleus of the lateral lemniscus. The latter region included a population of larger neurons (340-540 micron2) and had different connections with the inferior colliculus. The distribution of axonal labeling after injections in the nucleus sagulum was contrasted with the distribution of projections from several neighboring areas of the lateral tegmentum, including the dorsal nucleus of the lateral lemniscus. None of these areas exhibited connections with the superficial layers of the caudal cortex of the inferior colliculus, which was the major target in the inferior colliculus of the nucleus sagulum. Thus, the results indicated that the nucleus sagulum is distinguished from adjacent regions of the lateral tegmentum by its connectivity. Its association with midbrain auditory pathways is supported by these connections as well as ascending ones to the auditory thalamus.     

Connections among functional areas in the mustached bat auditory cortex

Connections among functional areas in the mustached bat's auditory cortex were examined by placing anatomical tracers in physiologically defined locations. We identified at least two and probably three channels connecting the various areas. One channel is formed by interconnections among areas containing neurons sensitive to frequency-modulated components (FMs) of the pulse and echo. These neurons are tuned to echo delay, a cue for target range, and thus define a ranging channel. An additional one or two channels are formed by interconnections among areas that contain neurons sensitive to the constant frequency components (CFs) of echoes. These neurons are of two main types: either sensitive to CFs of both pulse and echo (CF/CF neurons) or sensitive to a pulse FM and echo CF (FM-CF neurons). There was only a weak connection between the largest area of each type, suggesting they lie in different channels. Connections among areas in the ranging channel and echo CF-sensitive channel(s) were weak. Thus, the interconnections among functional areas in the mustached bat's auditory cortex define parallel channels for processing different types of biosonar information. Most corticocortical connections were patchy, in a manner suggestive of a columnar organization. The average width of the patches was approximately 360 μm. Based on the sizes of the functional areas, we estimate the auditory cortex contains a total of approximately 150 columns. Individual areas contain from as many as approximately 20 to as few as 1–4 columns. Each area had abundant projections outside of the auditory cortex. Connections within the cortex included the frontal, anterior cingulate, retrosplenial and perirhinal cortices, and the claustrum. Subcortical targets included the amygdyla, auditory thalamus, pons, pretectum, superior and inferior colliculi, and central gray. Projections within the cortex were of modest strength compared with several of the subcortical projections. Thus, the auditory areas themselves are the primary source of cortically processed biosonar information to the rest of the brain. J. Comp. Neurol. 391:366–396, 1998. © 1998 Wiley-Liss, Inc.     

Nuclei of the lateral lemniscus project directly to the thalamic auditory nuclei in the pigeon

Non-mesencephalic origins of ascending afferents to the thalamic auditory nuclei, nuclei ovoidalis (Ov) and semilunaris parovoidalis (SPO), were identified in the pigeon in retrograde tracing experiments. These origins comprise dorsal and ventral lateral lemniscal nuclei. Injections of anterograde tracers into these nuclei produced terminal labelling in SPO in particular. These experiments show for the first time that the inferior colliculus is not an obligatory relay to the auditory thalamus.     

Prefrontal connections of the parabelt auditory cortex in macaque monkeys

In the present study, we determined connections of three newly defined regions of auditory cortex with regions of the frontal lobe, and how two of these regions in the frontal lobe interconnect and connect to other portions of frontal cortex and the temporal lobe in macaque monkeys. We conceptualize auditory cortex as including a core of primary areas, a surrounding belt of auditory areas, a lateral parabelt of two divisions, and adjoining regions of temporal cortex with parabelt connections. Injections of several different fluorescent tracers and wheat germ agglutinin conjugated to horseradish peroxidase (WGA–HRP) were placed in caudal (CPB) and rostral (RPB) divisions of the parabelt, and in cortex of the superior temporal gyrus rostral to the parabelt with parabelt connections (STGr). Injections were also placed in two regions of the frontal lobe that were labeled by a parabelt injection in the same case. The results lead to several major conclusions. First, CPB injections label many neurons in dorsal prearcuate cortex in the region of the frontal eye field and neurons in dorsal prefrontal cortex of the principal sulcus, but few or no neurons in orbitofrontal cortex. Fine-grain label in these same regions as a result of a WGA–HRP injection suggests that the connections are reciprocal. Second, RPB injections label overlapping prearcuate and principal sulcus locations, as well as more rostral cortex of the principal sulcus, and several locations in orbitofrontal cortex. Third, STGr injections label locations in orbitofrontal cortex, some of which overlap those of RPB injections, but not prearcuate or principal sulcus locations. Fourth, injections in prearcuate and principal sulcus locations labeled by a CPB injection labeled neurons in CPB and RPB, with little involvement of the auditory belt and no involvement of the core. In addition, the results indicated that the two frontal lobe regions are densely interconnected. They also connect with largely separate regions of the frontal pole and more medial premotor and dorsal prefrontal cortex, but not with the extensive orbitofrontal region which has RPB and STGr connections. The results suggest that both RPB and CPB provide the major auditory connections with the region related to directing eye movements towards stimuli of interest, and the dorsal prefrontal cortex for working memory. Other auditory connections to these regions of the frontal lobe appear to be minor. RPB has connections with orbitofrontal cortex, important in psychosocial and emotional functions, while STGr primarily connects with orbital and polar prefrontal cortex.     

Topographic organization of neurons in the acoustic thalamus that project to the amygdala

Projections from the posterior thalamus to the amygdala have been implicated in the processing of the emotional significance of acoustic stimuli. The aim of the present studies was to determine which areas of the amygdala receive afferents from posterior thalamic structures that, in turn, receive afferents (presumably acoustic afferents) from the inferior colliculus. Projections from the posterior thalamus to the amygdala and striatum were examined in rats using anterograde and retrograde axonal transport techniques. Following injections of WGA-HRP into the posterior thalamic areas [including the medial division of the medial geniculate body, the posterior intralaminar nucleus (PIN) and the medial posterior complex (POM)], anterograde transport was seen in the lateral (AL), central (ACE), medial (AM), and basomedial (ABM) nuclei of the amygdala and in the amygdalostriatal transition area (AST) and posterior caudate putamen (CPU). Injection of WGA-HRP into each anterogradely labeled area produced retrograde transport to the posterior thalamus, but the pattern of transport varied with the site of the injection. Injections in AL and AST produced retrograde transport to neurons in the medial division of the medial geniculate body (MGM), PIN, suprageniculate nucleus (SG) and, to a lesser extent, the lateral posterior nucleus (LP). Injections of the ACE, AM, and ABM, in contrast, only labeled cells in POM. While the MGM, PIN, and SG each receive afferents from the inferior colliculus, POM does not. AL and AST, therefore, receive inputs from thalamic areas that, in turn, receive inputs from the inferior colliculus.     

Cortical projections to the superior colliculus in tree shrews (Tupaia belangeri)

The visuomotor functions of the superior colliculus depend not only on direct inputs from the retina, but also on inputs from neocortex. As mammals vary in the areal organization of neocortex, and in the organization of the number of visual and visuomotor areas, patterns of corticotectal projections vary. Primates in particular have a large number of visual areas projecting to the superior colliculus. As tree shrews are close relatives of primates, and they are also highly visual, we studied the distribution of cortical neurons projecting to the superior colliculus by injecting anatomical tracers into the colliculus. Since projections from visuotopically organized visual areas are expected to match the visuotopy of the superior colliculus, injections at different retinotopic locations in the superior colliculus provide information about the locations and organization of topographic areas in extrastriate cortex. Small injections in the superior colliculus labeled neurons in locations within areas 17 (V1) and 18 (V2) that are consistent with the known topography of these areas and the superior colliculus. In addition, the separate locations of clusters of labeled cells in temporal visual cortex provide evidence for five or more topographically organized areas. Injections that included deeper layers of the superior colliculus also labeled neurons in medial frontal cortex, likely in premotor cortex. Only occasional labeled neurons were observed in somatosensory or auditory cortex. Regardless of tracer injection location, we found that, unlike primates, a substantial projection to the superior colliculus from posterior parietal cortex is not a characteristic of tree shrews. J. Comp. Neurol. 521:1614–1632, 2013. © 2012 Wiley Periodicals, Inc.     

Central acoustic tract in an echolocating bat: an extralemniscal auditory pathway to the thalamus

To determine the sources and targets of auditory pathways that bypass the inferior colliculus in the mustache bat, we injected WGA-HRP in the medial geniculate body and related auditory nuclei of the thalamus as well as in the lower brainstem. We used electrophysiological methods to verify that the injection electrode was in an area responsive to sound. The only thalamic injections that produced retrograde transport to cells in auditory nuclei caudal to the inferior colliculus were those that included the suprageniculate nucleus. These injections labeled a group of large multipolar cells lying between the ventral nucleus of the lateral lemniscus and the superior olivary complex. Neurons in this cell group have also been shown to project to the deep layers of the superior colliculus in the mustache bat. The pathway revealed by these studies is almost identical to the "central acoustic tract" in which fibers course medial to the lateral lemniscus and bypass the inferior colliculus to reach the deep superior colliculus and the suprageniculate nucleus.     

Neural processing of target distance by echolocating bats: functional roles of the auditory midbrain

Using their biological sonar, bats estimate distance to avoid obstacles and capture moving prey. The primary distance cue is the delay between the bat's emitted echolocation pulse and the return of an echo. The mustached bat's auditory midbrain (inferior colliculus, IC) is crucial to the analysis of pulse-echo delay. IC neurons are selective for certain delays between frequency modulated (FM) elements of the pulse and echo. One role of the IC is to create these “delay-tuned”, “FM-FM” response properties through a series of spectro-temporal integrative interactions. A second major role of the midbrain is to project target distance information to many parts of the brain. Pathways through auditory thalamus undergo radical reorganization to create highly ordered maps of pulse-echo delay in auditory cortex, likely contributing to perceptual features of target distance analysis. FM-FM neurons in IC also project strongly to pre-motor centers including the pretectum and the pontine nuclei. These pathways may contribute to rapid adjustments in flight, body position, and sonar vocalizations that occur as a bat closes in on a target.     

Projections of auditory cortex to the medial geniculate body of the cat

The corticofugal projection from 12 auditory cortical fields onto the medial geniculate body was investigated in adult cats by using wheat germ agglutinin conjugated to horseradish peroxidase or biotinylated dextran amines. The chief goals were to determine the degree of divergence from single cortical fields, the pattern of convergence from several fields onto a single nucleus, the extent of reciprocal relations between corticothalamic and thalamocortical connections, and to contrast and compare the patterns of auditory corticogeniculate projections with corticofugal input to the inferior colliculus. The main findings were that (1) single areas showed a wide range of divergence, projecting to as few as 5, and to as many as 15, thalamic nuclei; (2) most nuclei received projections from approximately five cortical areas, whereas others were the target of as few as three areas; (3) there was global corticothalamic-thalamocortical reciprocity in every experiment, and there were also significant instances of nonreciprocal projections, with the corticothalamic input often more extensive; (4) the corticothalamic projection was far stronger and more divergent than the corticocollicular projection from the same areas, suggesting that the thalamus and the inferior colliculus receive differential degrees of corticofugal control; (5) cochleotopically organized areas had fewer corticothalamic projections than fields in which tonotopy was not a primary feature; and (6) all corticothalamic projections were topographic, focal, and clustered, indicating that areas with limited cochleotopic organization still have some internal spatial arrangement. The areas with the most divergent corticothalamic projections were polysensory regions in the posterior ectosylvian gyrus. The projection patterns were indistinguishable for the two tracers. These findings suggest that every auditory thalamic nucleus is under some degree of descending control. Many of the projections preserve the relations between cochleotopically organized thalamic and auditory areas, and suggest topographic relations between nontonotopic areas and nuclei. The collective size of the corticothalamic system suggests that both lemniscal and extralemniscal auditory thalamic nuclei receive significant corticofugal input.     

Thalamic connections of the auditory cortex in marmoset monkeys: core and medial belt regions

In this study and its companion, the cortical and subcortical connections of the medial belt region of the marmoset monkey auditory cortex were compared with the core region. The main objective was to document anatomical features that account for functional differences observed between areas. Injections of retrograde and bi-directional anatomical tracers targeted two core areas (A1 and R), and two medial belt areas (rostromedial [RM] and caudomedial [CM]). Topographically distinct patterns of connections were revealed among subdivisions of the medial geniculate complex (MGC) and multisensory thalamic nuclei, including the suprageniculate (Sg), limitans (Lim), medial pulvinar (PM), and posterior nucleus (Po). The dominant thalamic projection to the CM was the anterior dorsal division (MGad) of the MGC, whereas the posterior dorsal division (MGpd) targeted RM. CM also had substantial input from multisensory nuclei, especially the magnocellular division (MGm) of the MGC. RM had weak multisensory connections. Corticotectal projections of both RM and CM targeted the dorsomedial quadrant of the inferior colliculus, whereas the CM projection also included a pericentral extension around the ventromedial and lateral portion of the central nucleus. Areas A1 and R were characterized by focal topographic connections within the ventral division (MGv) of the MGC, reflecting the tonotopic organization of both core areas. The results indicate that parallel subcortical pathways target the core and medial belt regions and that RM and CM represent functionally distinct areas within the medial belt auditory cortex.     

The origin of the spinomesencephalic tract in the rat: An anatomical study using the retrograde transport of horseradish peroxidase

Projections of the middle temporal visual area, MT, and of visual cortex adjoining MT were investigated with autoradiographic methods in the prosimian primate, Galago senegalensis. Ipsilateral cortical targets of MT included area 17, area 18, cortex caudal to MT, cortex ventral to MT, and parietal-occipital cortex dorsal to MT. This pattern of projections suggests that extrastriate cortex contains a number of visual subdivisions in addition to MT. Contralateral projections were to MT and parietal-occipital cortex. Projections from MT to areas 17 and 18 connected regions representing similar parts of the visual hemifield while the location of callosal projections in MT matched the location of the injection site in the other hemisphere. Label in area 17 wac concentrated in layers I, III, and VI whereas other cortical areas were most densely labeled in the granular and supragranular layers. Subcortical projections of MT included the reticular nucleus of the thalamus, the lateral posterior nucleus, the superior pulvinar, the inferior pulvinar, the superior colliculus, and the pontine nuclei. The projection pattern to the superior and inferior pulvinar nuclei suggests that MT projects in a topographic manner to two subdivisions within each of these structures. Injections in cortex just outside of MT labeled area 18, inferotemporal cortex, parietal-occipital cortex, and, to a lesser extent, MT. The projections to inferotemporal cortex clearly distinguish the bordering cortex from MT. Contralateral cortical terminations were in locations corresponding to the injection site. Subcortical targets were generally similar to those seen after MT injections, although additional projections were observed depending on the location of the injection. Comparison of these results from the prosimian galago with studies in New and Old World monkeys indicates there are substantial similarities in projections. Thus, some of the cortical and thalamic subdivisions described for monkeys appear to exist in prosimians.     

Projections from cortex to tectum in the tree shrew, Tupaia glis.

Sensory neocortex of the tree shrew was divided into three main areas: the visual field, the auditory field, and the somatic field which includes motor cortex. Cortical cells which project to the tectum were identified by injecting HRP into superficial or deep layers of the superior colliculus and into various parts of the inferior colliculus. The main result is that these descending projections are well organized according to their origin in the three main sensory fields of the cortex. (1) Auditory field: labeled cells are found only in the core or auditory koniocortex, after injections of HRP in the pericentral area of the inferior colliculus; labeled cells are found in auditory belt areas after injections in posterior parts of the intermediate and deep layers of the superior colliculus, adjacent to the inferior colliculus. (2) Somatic field: labeled cells are also found in the somatic field after injections in the intermediate and deep layers of the superior colliculus, so that auditory and somatic fields probably overlap to some extent. The results do not exclude the possibility that somatic koniocortex has an exclusive target in the intermediate or deep layers of the superior colliculus. (3) Visual field: labeled cells are found only in the core or striate cortex after injections in the superficial layers of the superior colliculus. Labeled cells are found in the visual belt after injections in the rostral parts of the intermediate layers of the superior colliculus. When these results are related to ascending sensory pathways a picture emerges of a series of circuits or loops which interconnect parallel sensory pathways. These loops eventually reach the deep layers of the superior colliculus which of course have indirect access to motor neurons.     

Direct synaptic connections of axons from superior colliculus with identified thalamo-amygdaloid projection neurons in the rat: Possible substrates of a subcortical visual pathway to the amygdala

The aim of the present study was to identify synaptic contacts from axons originating in the superior colliculus with thalamic neurons projecting to the lateral nucleus of the amygdala. Axons from the superior colliculus were traced with the anterograde tracers Phaseolus vulgaris leucoagglutinin or the biotinylated and fluorescent dextran amine “Miniruby.” Thalamo-amygdaloid projection neurons were identified with the retrograde tracer Fluoro-Gold. Injections of Fluoro-Gold into the lateral nucleus of the amygdala labeled neurons in nuclei of the posterior thalamus which surround the medial geniculate body, viz. the suprageniculate nucleus, the medial division of the medial geniculate body, the posterior intralaminar nucleus, and the peripeduncular nucleus. Anterogradely labeled axons from the superior colliculus terminated in the same regions of the thalamus. Tecto-thalamic axons originating from superficial collicular layers were found predominantly in the suprageniculate nucleus, whereas axons from deep collicular layers were detected in equal density in all thalamic nuclei surrounding the medial geniculate body. Double-labeling experiments revealed an overlap of projection areas in the above-mentioned thalamic nuclei. Electron microscopy of areas of overlap confirmed synaptic contacts of anterogradely labeled presynaptic profiles originating in the superficial layers of the superior colliculus with retrogradely labeled postsynaptic profiles of thalamo-amygdaloid projection neurons. These connections may represent a subcortical pathway for visual information transfer to the amygdala. J. Comp. Neurol. 403:158–170, 1999. © 1999 Wiley-Liss, Inc.     

Spatial distribution of thalamic projections to the supplementary motor area and the primary motor cortex: A retrograde multiple labeling study in the macaque monkey

The exact knowledge on spatial organization of information sources from the thalamus to the supplementary motor area (SMA) and to the primary motor cortex (MI) has not been established. We investigated the distribution of thalamocortical neurons projecting to forelimb representations of the SMA and the MI using a multiple retrograde labeling technique in the monkey. The forelimb area of the SMA, and the distal and proximal forelimb areas of the MI were identified by electrophysiological techniques of intracortical microstimulation and single neuron recording. Injections were made into these three representations with three different dyes in the same animal (horseradish peroxidase conjugated to wheat germ agglutinin, diamidino yellow, and fast blue), and the thalamic neurons were retrogradely labeled. Injections into the SMA densely labeled thalamic neurons in nuclei ventralis lateralis pars oralis (VLo), ventralis lateralis pars medialis (VLm) and ventralis lateralis pars caudalis (VLc), but not in nucleus ventralis posterior lateralis pars oralis (VPLo). Injections into the MI labeled thalamic neurons primarily in VLo, VLc, and VPLo. We found that the distribution of projection neurons to the three areas was largely separate in the thalamus. However, in the middle part of VLo, and in a limited portion of VLc, thalamic neurons projecting to the SMA partially overlapped with those to the distal forelimb area of the MI. They overlapped little with those to the proximal forelimb area of the MI. We noted no overlap between the distributions of thalamic projection neurons to the distal and proximal forelimb areas of the MI. These findings suggest that the SMA and MI receive separate information from the thalamus, while sharing minor sources of common inputs. © 1995 Wiley-Liss, Inc.     

Role of commissural projections in the representation of bilateral auditory space in the barn owl's inferior colliculus

The central nucleus of the barn owl's inferior colliculus (ICc) contains a representation of both the ipsilateral and contralateral auditory hemifields. The representation of ipsilateral space is found in the "core" of the ICc, a subdivision defined by the terminal field of nucleus laminaris, the avian analogue of the medial superior olivary nucleus. The representation of contralateral space is found in the lateral portion of the "shell" of the ICc. The shell surrounds the core and is defined by the terminal field of the nucleus angularis, one of the cochlear nuclei. The representation of ipsilateral space in the core of the ICc may be accounted for by the crossed projection from the nucleus laminaris because most of the nucleus laminaris is devoted to a representation of contralateral space. We present evidence to suggest that the representation of contralateral space is due to a commissural projection from the core of one side to the lateral shell of the opposite side. Injection of horseradish peroxidase (HRP) into the lateral portion of the ICc shell produced retrogradely labeled somata in the core of the opposite side. Injection of tritiated proline into the core produced anterograde label confined to the lateral shell, thus confirming the observations made with HRP. Thus, for example, the left ICc core, which contains predominantly a representation of the left hemifield, innervates the right lateral shell, endowing it with a representation of the left, or contralateral hemifield. The representation of contralateral space in the lateral shell is ultimately conveyed to the external nucleus of the inferior colliculus where it contributes the horizontal axis to a two-dimensional map of space.     

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

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

Cells in the posterior thalamus project to both amygdala and temporal cortex: a quantitative retrograde double-labeling study in the rat

Auditory information from the posterior thalamus reaches the lateral nucleus of the amygdala (LA) by way of two pathways: a direct thalamo-amygdala projection and a polysynaptic thalamo-cortico-amygdala projection. However, the quantitative extent of thalamic neurons that project to the LA or to the auditory association cortex (AAC) is not known. Furthermore, the extent and topographical distribution of bifurcating cells that project to both LA and AAC are also unknown. Therefore, separate tracers were injected into LA and either into all of AAC or within discrete regions of AAC, such as temporal areas TE3 or perirhinal cortex (PRh), and quantitative analyses were performed on labeling within the subregions of the auditory thalamus in rats. Following LA injections, retrogradely labeled cells were most numerous in the posterior intralaminar nucleus (PIN; 48.0% of all labeled thalamic cells), whereas labeled cells following injections of the entire AAC were most numerous in the dorsal division of the medial geniculate nucleus (MGd; 32.9% of all labeled thalamic cells). Following AAC injections localized to only TE3, the MGd again had the majority of labeled cells (35.9%), whereas following AAC injections localized to PRh, the PIN had the most labeled cells (32.8%). Double-labeled cells were found in all the thalamic regions studied and were most commonly observed in the PIN (43.7% of all double-labeled cells following injections into LA and throughout the AAC). The percentage of double-labeled cells as a proportion of either LA-projecting or AAC-projecting cells varied among the thalamic nuclei studied, ranging from 2.9% up to 42.4%. The topographic distribution of double-labeled cells in the thalamic nuclei resembled that of single-labeled cells following LA injections more than single-labeled cells following AAC injection. These findings suggest that double-labeled cells contribute substantially to many of the direct thalamo-amygdala and indirect thalamo-AAC-amygdala projections. Among other functions, these bifurcating cells may help regulate the processing of input to the LA arriving from these two pathways to allow for certain types of plasticity in the LA during fear conditioning.     

Differential ascending projections to aural regions in the 60 kHz contour of the mustache bat's inferior colliculus

The inferior colliculus of the mustache bat is similar in many respects to the inferior colliculus of more commonly studied mammals. However, the isofrequency contour devoted to processing 60 kHz, the dorsoposterior division (DPD) is greatly expanded, encompassing an area approximately equal to one-third of the central nucleus. Of particular significance is that monaural and binaural neurons are segregated in the DPD into 4 spatially distinct aural regions. In this study we exploit the great enlargement of the 60 kHz region in the central nucleus of the inferior colliculus (ICc) of the mustache bat to determine the source of ascending projections to the 4 different aural regions of the DPD. Small iontophoretic deposits of HRP were made within each of the physiologically defined aural regions, and the locations and numbers of retrogradely labeled cells in the auditory brainstem nuclei were determined. Two major features of collicular organization were found. The first is that each aural region receives a unique set of projections from a subset of lower auditory nuclei and thus is distinguished both by its neural response properties and by the pattern of ascending projections it receives. The dorsomedial EE region receives inputs primarily from the ipsilateral intermediate nucleus of the lateral lemniscus (INLL) and ventral nucleus of the lateral lemniscus (VNLL), and the contralateral ICc. In contrast, the ventrolateral EE region receives projections from the ipsilateral medial superior olivary nucleus (MSO), VNLL, and INLL. The inputs to the EI region originate primarily from the dorsal nucleus of the lateral lemniscus (DNLL) and lateral superior olivary nucleus (LSO) bilaterally and from the ipsilateral INLL. The afferents to the EO region include the contralateral cochlear nucleus, the ipsilateral VNLL and INLL and MSO. The second major organizational feature is that the binaural nuclei of the brain-stem project upon the DPD in a more restricted manner than do some of the lower monaural nuclei, such as the VNLL and INLL, which project in a more widespread manner. The unique set of projections terminating in each aural region of the DPD suggests that the neurons should have substantially different properties, even when neurons in different regions are of the same general aural type. Moreover, the elucidation of the micro-organization of the DPD provides insights into the different ways that binaural properties of DPD neurons are created by the convergence of inputs from particular subsets of lower auditory nuclei.(ABSTRACT TRUNCATED AT 400 WORDS)     

Auditory cortical projections to the cat inferior colliculus

The projection from 11 auditory cortical areas onto the subdivisions of the inferior colliculus was studied in adult cats by using two different anterograde tracers to label corticocollicular (CC) axon terminals. The main results were that: 1) a significant CC projection arose from every field; 2) the principal inferior collicular targets were the dorsal cortex, lateral nucleus, caudal cortex, and intercollicular tegmentum, with only a sparse projection to the central nucleus; 3) the input was usually bilateral, with the ipsilateral side by far the most heavily labeled, and the contralateral projection was a symmetrical subset of the ipsilateral input; 4) the CC system is both divergent and convergent, with single cortical areas projecting to six or more collicular subdivisions, and each auditory midbrain subdivision receiving a convergent projection from two to ten cortical areas; 5) cortical areas devoid of tonotopic organization have topographic projections to collicular target nuclei; 6) the heaviest CC projection terminated in the caudal half of the inferior colliculus; and finally, 7) the relative strength of the corticocollicular labeling was far less than that of the corresponding corticothalamic projection in the same experiments. The CC system is strategically placed to influence both descending and ascending pathways arising in the inferior colliculus. Nuclei that participate in the premotor system, like the inferior collicular subdivisions that project to the pons, receive substantial corticofugal input. Both the dorsal (pericentral) and the lateral (external) nuclei of the inferior colliculus project to parts of the medial geniculate body whose closest auditory affiliations are with nontonotopic cortical regions involved in higher order auditory perception. The corticocollicular system may link brainstem and colliculothalamic circuits to coordinate premotor and perceptual aspects of hearing. J. Comp. Neurol. 400:147–174, 1998. © 1998 Wiley-Liss, Inc.