GAD- and GABA-immunoreactivity in the ascending auditory pathway of horseshoe and mustached bats.

A comparative study of the immunostain to antibodies directed against glutamic acid decarboxylase (GAD) and gamma-aminobutyric acid (GABA) in the ascending auditory pathway was carried out in horseshoe bats (Rhinolophus rouxi) and mustached bats (Pteronotus parnellii). In both species GAD/GABA-positive puncta (presumed axonal boutons) and GAD/GABA-positive cells were found in the cochlear nucleus, the superior olivary complex, the nuclei of the lateral lemniscus the inferior colliculus, and the medial geniculate body. General features of the immunostaining pattern in the auditory pathway agree with observations in other mammals. Quantitative analysis of puncta distribution shows that many auditory centers are characterized by subregional differences in puncta density and distribution. This indicates local differences in putatively inhibitory input related to connectivity and tonotopic organization. The following species characteristic features were found: 1) The dorsal non-laminated portion of the dorsal cochlear nucleus in horseshoe bats lacks the GAD/GABA-immunoreactive cells typical for the ventral laminated portion and the dorsal cochlear nucleus of other species. Clearly, a cytoarchitectonic specialization is accompanied by a loss of putatively GABAergic local inhibitory circuits. 2) The ventral division of the medial geniculate body of the mustached bat lacks GAD/GABA-immunopositive cells. Such cells are present in the horseshoe bat and other mammals. This finding implies functional differences in the organization of the medial geniculate body within the same mammalian order.     

An extralemniscal component of the mustached bat inferior colliculus selective for direction and rate of linear frequency modulations

Frequency modulations (FMs) are prevalent in human speech, and are important acoustic cues for the categorical discrimination of phonetic contrasts. For bats, FM sweeps are also important for communication and are often the only component in echolocation calls. Auditory neurons tuned to the direction and rate of FM might underlie the encoding of rapid frequency transitions. In the mustached bat, we have discovered a population of such FM selective cells in an area interposed between the central nucleus of the inferior colliculus (ICC) and the nuclei of the lateral lemniscus (NLL). We believe this area to be the ventral extent of the external nucleus of the inferior colliculus (ICXv). To describe FM selectivity of neurons in the ICXv and to compare it to other midbrain nuclei, up- and down-sweeping linear FM stimuli were presented at different modulation rates. Extracellular recordings were made from 171 single units in the ICC, ICXv, and NLL of 10 mustached bats. In the ICXv, there was a much higher degree of FM selectivity than in ICC or NLL and a consistent preference for upward over downward FM sweeps. Anterograde and retrograde transport was examined following focal injections of wheatgerm agglutinin-horseradish peroxidase (WGA-HRP) into ICXv. The main targets of anterograde transport were the deep layers of the superior colliculus and the suprageniculate division of the medial geniculate body. The primary site of retrograde transport was the nucleus of the central acoustic tract in the brainstem. Thus, the ICXv appears to be a part of the central acoustic tract, an extralemniscal pathway linking the auditory brainstem directly to a multimodal nucleus of the thalamus.     

Electron microscopy of classically stained astrocytes

The ascending connections of the lateral lemniscus were studied in the cat and squirrel monkey (Saimiri sciureus). In both species, the central nucleus of the inferior colliculus receives a massive projection from the lateral lemniscus. Only a few lemniscal fibers were found to terminate in the external nucleus of the inferior colliculus. The commissure of the lateral lemniscus originates from the dorsal nucleus of the lateral lemniscus and projects to the contralateral dorsal nucleus and the contralateral central nucleus of the inferior colliculus. No lemniscal fibers were seen ascending in the inferior brachium or terminating in the principal division of the medial geniculate body. A bundle of fibers was observed, however, which passed medial to the inferior brachium and terminated in the magnocellular or internal division of the medial geniculate. The bundle degenerated after lesions confined to the lateral lemniscus and is probably identical with the central acoustic tract of earlier workers. Evidence is presented that the fibers of the bundle are of spinal origin. Since the lemniscal fibers which ascend to the thalamus appear to be non-auditory, it is suggested that the inferior colliculus is an obligatory relay station in the classical auditory system and that the inferior brachium is the only ascending pathway of the system to project upon the thalamus.     

Cytoarchitectonic and connectional organization of the ventral lateral geniculate nucleus in the cat

The ventral lateral geniculate nucleus is a small extrageniculate visual structure that has a complex cytoarchitecture and diverse connections. In addition to small-celled medial and lateral divisions, we cytoarchitectonically defined a small-celled dorsal division. A large-celled intermediate division intercalated between the three small-celled divisions, which we divided into medial and lateral intermediate subdivisions. In WGA-HRP injection experiments, the different cytoarchitectonic divisions were shown to have connections with different nuclei. The medial division was reciprocally connected to the pretectum and projected to the superficial layers of the superior colliculus and the intralaminar nuclei. The medial intermediate division received projections from the intermediate layer of the superior colliculus and the lateral and interpositus posterior cerebellar nuclei, and projected to the intermediate layer of the superior colliculus, the periaqueductal gray of midbrain, and the intralaminar nuclei. The lateral intermediate divisions received projections from the pretectum, the intermediate layer of the superior colliculus, and the lateral and interpositus posterior cerebellar nuclei, and projected to the pretectum, superficial layers of the superior colliculus, and the pulvinar. The lateral division received projections from superficial layers of the superior colliculus and had reciprocal connections with the pretectum. The dorsal division received projections from the pretectum and had reciprocal connections with the periaqueductal gray of midbrain. The different cytoarchitectonic divisions of the ventral lateral geniculate nucleus are thus suggested to play different functional roles related to vision, eye and head movements, attention, and defensive reactions.     

Origin of ascending projections to inferior colliculus in the mustache bat,Pteronotus parnellii

The origins of pathways to the inferior colliculus of the mustache bat were identified by retrograde transport of horseradish peroxidase (HRP). A specific goal of this study was to obtain evidence that would help determine whether the nuclei, shown in the previous paper to have unusual cytoarchitectural features, are unique to bats, or whether they are homologous to areas that are not well differentiated in other mammals. The auditory pathways in the lower brain stem of Pteronotus appear to conform to the same basic organization as in other mammals: After injection of HRP into one inferior colliculus, labeled cells are located contralaterally in the cochlear nucleus, ipsilaterally in the medial superior olive, bilaterally in the lateral superior olive, ipsilaterally in the ventral and intermediate nuclei of the lateral lemniscus, and bilaterally in the dorsal nucleus of the lateral lemniscus. These patterns of labeling provide a basis for understanding how the specialized auditory areas of the bat may be organized within a basic plan of mammalian auditory systems. In the anteroventral cochlear nucleus the unusually small spherical cells seem to be homologous to stellate cells in the anteroventral cochlear nucleus of the cat. In the superior olive, differences in patterns of labeled cells distinguish the medial from the lateral superior olive. In the lateral lemniscus the pattern of labeled cells shows clear differences between the two special parts, intermediate and ventral nuclei, as well as between these and the dorsal nucleus of the lateral lemniscus. The results are consistent with the hypothesis that the unusual auditory nuclei of the bat have homologues in mammals whose auditory systems are not specialized for echolocation.     

Acoustic chiasm. IV: Eight midbrain decussations of the auditory system in the cat.

Conventional retrograde and orthograde axonal transport tract-tracing techniques were used in cats to explore the auditory decussations and commissures in the upper pons and midbrain. In all, 8 decussations differing either in origin or in contralateral termination were found. Three of the 8 decussations (from the dorsal nucleus of the lateral lemniscus to the contralateral dorsal nucleus of the lateral lemniscus, from the dorsal nucleus of the lateral lemniscus to the contralateral inferior colliculus, from the sagulum to the contralateral sagulum) reach their targets via the commissure of Probst. The remaining 5 decussations (from the inferior colliculus to the contralateral inferior colliculus or medial geniculate, from the intermediate nucleus of the lateral lemniscus to the contralateral medial geniculate, from the sagulum to the contralateral inferior colliculus or medial geniculate) reach their targets via the commissure of the inferior colliculus. The results also suggest that the commissure of Probst is not a general avenue for decussating auditory fibers of the lateral lemniscus but is instead a specific avenue only for fibers from the dorsal nucleus of the lateral lemniscus and sagulum. The results also show that, in the cat at least, the dorsal nucleus of the lateral lemniscus does not project beyond the inferior colliculus to either the superior colliculus or medial geniculate--the cells previously reported as doing so are probably those of the immediate neighbors of the dorsal nucleus, the intermediate nucleus of the lateral lemniscus and sagulum.     

Topographical organization of the inferior collicular projection and other connections of the ventral nucleus of the lateral lemniscus in the cat

The topographic distribution of projections from the ventral nucleus of the lateral lemniscus (VNLL) in the cat was investigated with the autoradiographic tracing method. The origin of minor projections was verified by retrograde tracing methods. Small injections of tritiated leucine were placed in restricted zones of VNLL. A major afferent fiber system to the inferior colliculus was labeled in all cases. From the injection site labeled fibers coursed through and around the nuclei of the lateral lemniscus to enter the ipsilateral inferior colliculus. Regardless of the position or small size of the injection, labeled fibers distributed widely in the inferior colliculus. Fibers ended in the central nucleus and deeper layers of the dorsal cortex in most cases. There was also labeling in the ventrolateral nucleus, but very few fibers ended as lateral as the lateral nucleus. A small number of labeled fibers passed from the inferior colliculus into the nucleus of the brachium of the inferior colliculus and adjacent tegmental areas. Some labeled fibers entered the commissure of the inferior colliculus where they were traced into the dorsal cortex and rostral pole of the inferior colliculus on the side contralateral to the injection site. Though the projections labeled in individual cases were similar in their divergent pattern within the central nucleus of the inferior colliculus, specific variations in the pattern were found. The dorsal zone of VNLL projected more heavily to the deeper layers of the dorsal cortex and an adjacent field in the central nucleus than the other zones. Dorsal injections in the middle zone of VNLL, on the other hand, labeled the medial part of the central nucleus more heavily, whereas ventral injections in the middle zone resulted in heavier lateral labeling. The ventral zone of VNLL projected heavily to a central field in the central nucleus. In addition to this major afferent system of VNLL to the inferior colliculus, a smaller descending projection was found. The descending projection ended mainly in the dorsomedial periolivary region and ventral nucleus of the trapezoid body. However, in some cases a few fibers were traced to the cochlear nuclei. Finally, we observed projections to the medial geniculate body from the dorsal and ventral zones of VNLL that ended diffusely in the medial division of the medial geniculate body. Possibly some fibers from the dorsal zone contribute to a broader projection of the lateral tegmentum to the dorsal division of the medial geniculate body.     

The tecto-thalamic connections in the brain of the rabbit

We have correlated the tectal connections and cytoarchitecture of regions in the rabbit's midbrain and caudal thalamus. The inferior colliculus projects ipsilaterally to the central gray, superior colliculus, and via the brachium of the inferior colliculus to its interstitial nucleus and the parabrachial region of the midbrain tegmentum. From the brachium, fibers fan out to the principal and internal divisions of the medial geniculate. A smaller contralateral pathway sweeps into the contralateral inferior colliculus and in its brachium to the interstitial nucleus, the parabrachial region, and the internal and principal divisions of the medial geniculate. The superior collicular projection is mainly ipsilateral. Medially, fibers terminate in the central gray and pretectal area. Laterally, fibers ascend in the superior brachium to parabrachial region, suprageniculate pretectal nucleus, posterior complex, caudodorsal internal division of the medial geniculate, and to a discrete part of the ventral nucleus of later geniculate. A component of the commissure of Gudden originates in the rostral superior colliculus and terminates in the contralateral ventral lateral geniculate, posterior complex, pretectal area and midbrain tegmentum. Interconnections between the colliculi and overlap of their projections in the parabrachial region, the central gray, and the internal division of the medial geniculate are described.     

Projections from the cochlear nuclei in the mustache bat, Pteronotus parnellii

Ascending projections of the cochlear nuclei in the mustache bat were analyzed by anterograde transport of [3H]-leucine and by retrograde transport of HRP. We were particularly interested in pathways to two parts of the system: (1) to the medial superior olive, because this nucleus is missing in most echolocating bats, but appears to be present in the mustache bat, and (2) to the intermediate and ventral nuclei of the lateral lemniscus, because these nuclei are hypertrophied and highly differentiated in all echolocating bats that we have examined. The results show a highly systematic projection from the anteroventral cochlear nucleus to all of the auditory nuclei in the brain stem. After an injection of [3H]-leucine in the anterior and dorsal part of the anteroventral cochlear nucleus, presumably in a region sensitive to low frequencies, label is seen in the following locations: ipsilateral to the injection in the lateral part of the lateral superior olive; bilaterally in the dorsal part of the medial superior olive; contralateral to the injection in the dorsal parts of the intermediate and ventral nuclei of the lateral lemniscus; and in the anterolateral part of the central nucleus of the inferior colliculus. After an injection of [3H]-leucine in a posterior part of the anteroventral cochlear nucleus, presumably in a region sensitive to high frequencies, labeling is in the same set of nuclei, but within each nucleus the label is now in a different location: medially in the lateral superior olive, ventrally in the medial superior olive, ventrally in each division of the ventral and intermediate nuclei of the lateral lemniscus, and medially in the central nucleus of the inferior colliculus. Projections from the entire anteroventral cochlear nucleus to the inferior colliculus are confined to the ventral two-thirds of the central nucleus. The dorsal one-third of the central nucleus of the inferior colliculus is the principal target of the dorsal cochlear nucleus and may be a target of the posteroventral cochlear nucleus. Both of these nuclei appear to project sparsely to the ventral parts of the inferior colliculus. We conclude first that the bilateral input to the medial superior olive in the mustache bat is similar to the input seen in other mammals. Thus this bat has a neural structure which is associated with the analysis of binaural time differences and which usually is seen only in animals with heads large enough to create interaural time differences greater than those available to Pteronotus.(ABSTRACT TRUNCATED AT 400 WORDS)     

Inputs to combination-sensitive neurons of the inferior colliculus.

In the mustached bat, combination-sensitive neurons display integrative responses to combinations of acoustic elements in biosonar or social vocalizations. One type of combination-sensitive neuron responds to multiple harmonics of the frequency-modulated (FM) components in the sonar pulse and echo of the bat. These neurons, termed FM-FM neurons, are sensitive to the pulse-echo delay and may encode the distance of sonar targets. FM-FM neurons are common in high-frequency regions of the central nucleus of the inferior colliculus (ICC) and may be created there. If so, they must receive low-frequency inputs in addition to the expected high-frequency inputs. We placed single deposits of a tracer at FM-FM recording sites in the ICC and then analyzed retrograde labeling in the brainstem and midbrain. We were particularly interested in labeling patterns suggestive of low-frequency input to these FM-FM neurons. In most nuclei containing labeled cells, there was a single focus of labeling in regions thought to be responsive to high-frequency sounds. More complex labeling patterns were observed in three nuclei. In the anteroventral cochlear nucleus, labeling in the anterior and marginal cell divisions occurred in regions thought to respond to low-frequency sounds. This labeling comprised 6% of total brainstem labeled cells. Labeling in the intermediate nucleus of the lateral lemniscus and the magnocellular part of the ventral nucleus of the lateral lemniscus together comprised nearly 40% of all labeled cells. In both nuclei, multiple foci of labeling occurred. These different foci may represent groups of cells tuned to different frequency bands. Thus, one or more of these three nuclei may provide low-frequency input to high-frequency-sensitive cells in the ICC, creating FM-FM responses. We also examined whether ICC neurons responsive to lower frequencies project to high-frequency-sensitive ICC regions; only 0.15% of labeling originated from these lower frequency representations. If the spectral integration of FM-FM neurons is created at the level of the ICC, these results suggest that neurons of the anteroventral cochlear nucleus or monaural nuclei of the lateral lemniscus may provide the essential low-frequency input. In contrast, there is little evidence that the low-frequency representation of the ICC contributes to these integrative responses.     

Connectional basis for frequency representation in the nuclei of the lateral lemniscus of the bat Eptesicus fuscus

To study the role of the lateral lemniscus as a link in the ascending auditory pathway, injections of neuronal tracers were placed in the anteroventral cochlear nucleus (AVCN) and in the inferior colliculus of the bat Eptesicus fuscus. To correlate the anatomical results with tonotopic organization, the characteristic frequency of cells at each injection site was determined electrophysiologically. Pathways from AVCN diverge to 3 major targets in the lateral lemniscus, the intermediate nucleus and 2 divisions of the ventral nucleus (VNLL). Projections from these 3 nuclei then converge at the inferior colliculus. One cell group is particularly notable for its cytoarchitectural appearance. It is referred to here as the columnar area of VNLL because its cells are organized as a tightly packed matrix of columns and rows. The connections of the columnar area are organized in sheets that are precisely related to the tonotopic organization of both AVCN and the inferior colliculus. Sheets of cells in the dorsal part of the columnar area receive projections from low-frequency parts of AVCN and project to low-frequency parts of the inferior colliculus. These sheets of connections occupy successively more ventral locations as the tonotopic focus of the injection site increases in frequency. The entire range of frequencies audible to the bat is systematically represented along the dorsal-ventral dimension of the columnar area. Because each column is only 20-30 cells in height, frequency representation must be compressed in this dimension. Within the columnar area there is an overrepresentation of frequencies between 25 and 50 kHz, which corresponds roughly to the range of the FM echo-location call in Eptesicus. The connections of the other nuclei of the lateral lemniscus are not as precisely related to the tonotopic organization of the system as are those of the columnar area.     

Auditory pathways to the cortex in Tupaia glis

The auditory system of the tree shrew, Tupaia glis, was investigated by identifying axonal degeneration after lesions of the lateral lemniscus, the inferior colliculus, the medial geniculate nucleus and the auditory cortex. The results show that the lateral lemniscus projects to the central nucleus of the inferior colliculus which in turn projects principally to the ventral division of the medial geniculate nucleus but to a lesser extent to the magnocellular division of the medial geniculate nucleus. The final step in the pathway to the cortex is achieved by a projection from the ventral division to the fourth layer of auditory koniocortex. There appear to be several auditory pathways parallel to this primary path. The lateral lemniscus projects to the dorsal division of the medial geniculate nucleus; the deeper layers of the superior colliculus project to the posterior nucleus; and both the dorsal division and the posterior nucleus project to the belt caudal to auditory koniocortex. The caudal division of the medial geniculate nucleus may constitute a relay in still another path from the pericentral division of the inferior colliculus. Finally, the magnocellular division also appears to be distinct insofar as its cortical projections are confined chiefly to the deeper layers. A comparison between the tree shrew and the cat reveals a similar organization in the two species. In the cat the starting point for understanding the organization of the several auditory pathways is the distinction between a core cortical zone which corresponds to koniocortex and to AI and a peripheral belt. The core receives essential projections from the ventral division; the belt receives sustaining projections from the cell groups which surround the ventral division. It is reasonable to hypothesize that this difference between the core and the belt is characteristic of all mammals.     

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.     

The medial geniculate body of the tree shrew, Tupaia glis. I. Cytoarchitecture and midbrain connections.

In this study of the medial geniculate body in the tree shrew eight subdivisions are identified on the basis of differences recognized in Nissl-stained material. Experiments using the methods of anterograde and retrograde axonal transport and anterograde degeneration show that each subdivision has a unique pattern of connections with the midbrain. The ventral division of the medial geniculate body contains at least two subdivisions, the ventral nucleus and the caudomarginal nucleus. The ventral nucleus is characterized by densely-packed cells and receives topographically organized projections from the central nucleus of the inferior colliculus. The caudomarginal nucleus, on the other hand, receives its major midbrain projections from the medial nucleus in the inferior colliculus. In the dorsal division four subdivisions are distinguished. The suprageniculate nucleus contains large, loosely-packed cells and receives projections from the deep layers of the superior colliculus and from the midbrain tegmentum. The dorsal nucleus receives projections from the midbrain tegmentum. The deep dorsal and anterodorsal nuclei have neurons which resemble those in the dorsal nucleus. Both receive projections from the roof nucleus of the inferior colliculus but the deep dorsal nucleus receives an additional projection from the parabrachial tegmentum. The medial division has a rostral and a caudal subdivision. The ascending projections to the rostral nucleus are from the lateral zone in the inferior colliculus and from the spinal cord. The caudal nucleus contains cells with large somas and receives projections from most of the midbrain areas which project to the other subdivisions of the medial geniculate body.     

Second order auditory pathways in the chimpanzee

Substantial portions of the dorsal, and almost the entire posteroventral and anteroventral (Av) cochlear nuclei were aspirated unilaterally in a chimpanzee. Axonal degeneration was studied by the Fink-Heimer method. The greatest amount of degeneration was followed medially from the region of Av into the lateral part of the trapezoid body. Degeneration also coursed around the superior surface of the restiform body and was traced into the dorsal and intermediate acoustic striae. Within the superior olivary complex, degeneration was distributed to: the ipsilateral lateral superior olive; laterally and medially oriented dendrites of the ipsilateral and contralateral medial superior olivary nuclei respectively (some periosomatic degeneration also was present bilaterally); the contralateral medial trapezoid nucleus; retro-olivary and preolivary cell groups bilaterally. Abundunt degeneration passed into the contralateral lateral lemniscus and was distributed largely to its ventral nucleus. The contralateral central nucleus of the inferior colliculus was a major site of termination of ascending second order auditory fibers. The caudal tip of the ipsilateral ventral nucleus of the lateral lemniscus received abundant degeneration, but this diminished rostrally. The ipsilateral inferior colliculus contained a moderate amount of degeneration. A fair number of degenerated second order auditory fibers ascended in the contralateral brachium of the inferior colliculus and were distributed both to the principal and magnocellular divisions of the medial geniculate body. This pathway appears to represent a phylogenetic advance in the brain of the great ape.     

Functional organization of auditory cortex in the Mongolian gerbil (Meriones unguiculatus). IV. Connections with anatomically characterized subcortical structures

The subcortical connections of the four tonotopically organized fields of the auditory cortex of the Mongolian gerbil, namely the primary (AI), the anterior (AAF), the dorsoposterior (DP) and the ventroposterior field (VP), were studied predominantly by anterograde transport of biocytin injected into these fields. In order to allow the localization of connections with respect to subdivisions of subcortical auditory structures, their cyto-, fibre- and chemoarchitecture was characterized using staining methods for cell bodies, myelin and the calcium-binding protein parvalbumin. Each injected auditory cortical field has substantial and reciprocal connections with each of the three subdivision of the medial geniculate body (MGB), namely the ventral (MGv), dorsal (MGd) and medial division (MGm). However, the relative strengths of these connections vary: AI is predominantly connected with MGv, AAF with MGm and MGv, and DP and VP with MGd and MGv. The connections of at least AI and MGv are topographic: injections into caudal low-frequency AI label laterorostral portions of MGv, whereas injections into rostral high-frequency AI label mediocaudal portions of MGv. All investigated auditory fields send axons to the suprageniculate, posterior limitans, laterodorsal and lateral posterior thalamic nuclei, with strongest projections from DP and VP, as well as to the reticular and subgeniculate thalamic nuclei. AI, AAF, DP and VP project to all three subdivisions of the inferior colliculus, namely the dorsal cortex, external cortex and central nucleus ipsilaterally and to the dorsal and external cortex contralaterally. They also project to the deep and intermediate layers of the ipsilateral superior colliculus, with strongest projections from DP and VP to the lateral and basolateral amygdaloid nuclei, the caudate putamen, globus pallidus and the pontine nuclei. In addition, AAF and particularly DP and VP project to paralemniscal regions around the dorsal nucleus of the lateral lemniscus (DNLL), to the DNLL itself and to the rostroventral aspect of the superior olivary complex. Moreover, DP and VP send axons to the dorsal lateral geniculate nucleus. The differences with respect to the existence and/or relative strengths of subcortical connections of the examined auditory cortical fields suggest a somewhat different function of each of these fields in auditory processing.     

Convergence and divergence of ascending binaural and monaural pathways from the superior olives of the mustached bat

The lateral superior olive and medial superior olive give rise to pathways that terminate in the dorsal nucleus of the lateral lemniscus and central nucleus of the inferior colliculus. In most mammals, neurons in both the medial and lateral superior olives are binaural, but in the mustached bat most neurons in the medial superior olive are monaural. The aims of this study were to determine how the inputs to the medial superior olive contribute to its monaurality and to determine whether the ascending projections from the lateral and medial superior olives overlap or rema in segregated at their targets. Injections of two different tracers were placed in tonotopically matched areas of the lateral and medial superior olives in the same animal. Retrograde transport from injections in the medial superior olive labeled spherical cells in the contralateral anteroventral cochlear nucleus and principal cells in the ipsilateral medial nucleus of the trapezoid body. Few cells were labeled in ipsilateral cochlear nucleus. Anterograde transport resulted in tonotopically specific distributions of label with the same laterality as in nonecholocating mammals. In the dorsal nucleus of the lateral lemniscus, label from the lateral and medial superior olives largely overlapped. In the inferior colliculus, label from the two sources overlapped in the high and low frequency ranges, but in the frequency range around 60 kHz, label from the medial superior olive extended more dorsally than that from the lateral superior olive. These results indicate that projections of the lateral and medial superior olives overlap extensively at their targets. © 1995 Willy-Liss, Inc.     

Neurotransmitter-specific uptake and retrograde transport of [3H]glycine from the inferior colliculus by ipsilateral projections of the superior olivary complex and nuclei of the lateral lemniscus.

Neurotransmitter-specific uptake and retrograde axonal transport of [3H]glycine were used to identify glycinergic projections to the inferior colliculus in chinchillas and guinea pigs. Six h after injection of [3H]glycine in the inferior colliculus, autoradiographically labeled cells were found ipsilaterally in the ventral nucleus of the lateral lemniscus, the lateral superior olive and the dorsomedial periolivary nucleus. These 3 regions accounted for 95% of the labeled projection neurons, with the remainder scattered elsewhere in the ipsilateral superior olivary complex. No labeled cells were found contralaterally even after survival times as long as 24 h. Retrograde transport of HRP from the inferior colliculus in these same cases confirmed the presence of additional projections that did not accumulate [3H]glycine. These included ipsilateral projections from the medial superior olive and cochlear nucleus and contralateral projections from the inferior colliculus, dorsal nucleus of the lateral lemniscus, lateral superior olive, periolivary nuclei and cochlear nucleus. The results implicate uncrossed projections from the ventral nucleus of the lateral lemniscus, lateral superior olive, and dorsomedial periolivary nucleus as the principal sources of inhibitory glycinergic inputs to the inferior colliculus.     

Sources of subcortical projections to the superior colliculus in the cat

A comprehensive search for subcortical projections to the cat superior colliculus was conducted using the retrograde horseradish peroxidase (HRP) method. Over 40 different subcortical structures project to the superior colliculus. The more notable among these are grouped under the following categories. Visual structures: ventral lateral geniculate nucleus, parabigeminal nucleus, pretectal area (nucleus of the optic tract, posterior pretectal nucleus, nuclei of the posterior commissure). Auditory structures: inferior colliculus (external and pericentral nuclei), dorsomedial periolivary nucleus, nuclei of the trapezoid body, ventral nucleus of the lateral lemniscus. Somatosensory structures: sensory trigeminal complex (all divisions, but mainly the γ division of nucleus oralis), dorsal column nuclei (mostly cuneate nucleus), and the lateral cervical nucleus. Catecholamine nuclei: locus coeruleus, raphe dorsalis, and the parabrachial nuclei. Cerebellum: medial, interposed, and lateral nuclei, and the perihypoglossal nuclei. Reticular areas: zona incerta, substantia nigra, midbrain tegmentum, nucleus paragigantocellularis lateralis, and the hypothalamus. Evidence is presented that only the parabigeminal nucleus, the nucleus of the optic tract, and the posterior pretectal nucleus project to the superficial collicular layers (striatum griseum superficiale and stratum opticum), while all other afferents terminate in the deeper layers of the colliculus. Also presented is information concerning the rostrocaudal distribution of some of these afferent connections. These findings stress the multiplicity and diversity of inputs to the deeper collicular layers, and more specifically, identify multiple sources of the physiologically well-known representations of the somatic and auditory modalities in the colliculus.