Cross-modal innervation of primary visual cortex by auditory fibers in congenitally anophthalmic mice

Auditory-visual cross-modal innervation was examined in control (sighted, ZRDCT-N) and congenitally anophthalmic (eyeless, ZRDCT-AN) mice using electrophysiological recording and pathway tracing with carbocyanine dyes. Electrophysiological data demonstrate that the primary visual cortex of congenitally eyeless, blind, mice receives auditory stimuli. Neuroanatomical data demonstrate a direct connection between the inferior colliculus (IC) and visual cortex. Our experiments provide new information about how the brain adapts to the loss of sight.     

The thalamic projection to cortical area 17 in a congenitally anophthalmic mouse strain

The thalamocortical projection innervating area 17 in the congenitally anophthalmic ZRDCT-An mouse was compared with its counterpart in normal-eyed mice of different strains by the horseradish peroxidase method. The results indicate that, despite absence of retinal afferents and a reduction of its neuronal population to 76% of normal, the dorsal nucleus of the lateral geniculate body in the ZRDCT-An mouse projects to area 17 in an essentially normal topographic pattern. Our evidence also indicates the existence, both in normal-eyed and ZRDCT-An mice, of an extrageniculate thalamocortical projection to area 17, arising in particular from the nucleus lateralis posterior but in lesser degree also from the nucleus lateralis dorsalis. In both normal and anophthalmic mice, this extrageniculate projection favors the lateral and posterior parts of area 17; however, the projection to these parts is considerably greater in the congenitally anophthalmic than in the normal-eyed mouse.Compensatory innervation thus appears to occur in the anophthalmic mouse not only at the level of the primary retinorecipient nucleus of the geniculostriate system, but also at the level of the visual cortex that is once removed from the direct effects of congenital denervation.     

Trial sequence of changed unit activity in auditory system of alert rat during conditioned response acquisition and extinction.

1. The activity of units in the auditory system of alert, freely moving rats was studies during the acquisition and extinction of a tone-signaled, appetitive classically conditioned response. Responses of neurons in inferior colliculus (N = 28), medial geniculate (N = 32), posterior nucleus of thalamus (N = 28), pretectal region (N = 19), and cortex (N = 100) were studies in 74 rats across 10-trial blocks. 2. During behavioral acquisition, neurons in posterior nucleus of thalamus were the first to show response increments to CS+ onset. They were followed by neurons in cortex, pretectal region, medial geniculate, inferior colliculus and by movement behavior. 3. Prestimulus background rates during acquisition showed significant decrements in cortical neurons. These background decrements began to be evidenced in the trial series before the response increases in posterior nucleus. These data strengthened the suggestion of a previous study that posterior nucleus responses could be dependent on tonic modulation from cortex. 4. Extinction appeared to be largely a reverse of acquisition. Cortex and behavior showed response decrements first in the trial series. They were followed by medial geniculate, pretectal region, posterior nucleus, and inferior colliculus neurons. 5. The hypothesis was advanced that the auditory lemniscal adjunct afferent system may play a primary role in the early phases of auditory conditioned-response acquisition.     

Visual and auditory pathways contain cholecystokinin: evidence from immunofluorescence and retrograde tracing

The distribution of cholecystokinin (CCK) in the visual and auditory systems of the rat was studied with combined immunofluorescence and fluorescence retrograde tracing techniques. Double-labeled projection neurons in the pathway from the dorsal lateral geniculate nucleus to area 17 of visual cortex, and from the superior olive to the inferior colliculus demonstrated the presence of CCK-containing pathways in the ascending visual and auditory systems. Thus, CCK can be viewed as a neurotransmitter/neuromodulator candidate in ascending sensory systems.     

Immunocytochemical evidence for the involvement of glycine in sensory centers of the rat brain.

Glycine-like immunoreactivity was localized to a number of sites in the rat brain which are involved in processing sensory information. In the auditory and vestibular systems, glycine immunoreactivity was seen in dorsal and ventral cochlear nuclei, superior olive, trapezoid body, medial and lateral vestibular nuclei, and inferior colliculus. Staining in the visual system was seen in retina, dorsal lateral geniculate nucleus, and superior colliculus. The olfactory system exhibited staining in the olfactory bulb and accessory olfactory formation. Somatosensory centers with glycine immunoreactivity included the dorsal column nuclei, spinal trigeminal nucleus, principal sensory nucleus of V, reticular formation, and periaqueductal gray. Glycine-immunoreactive neurons were also seen in cerebellar cortex, deep cerebellar nuclei, hippocampus, cerebral cortex, and striatum. The distribution of staining indicates that glycine plays a major role in sensory centers with actions at both strychnine-sensitive and strychnine-insensitive receptors.     

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.     

Some topographical connections of the striate cortex with subcortical structures in Macaca fascicularis

Summary Subcortical connections of the striate cortex with the superior colliculus (SC), the lateral pulvinar (Pl), the inferior pulvinar (Pi) and the dorsal lateral geniculate nucleus (LG) were studied in the macaque monkey, Macaca fascicularis, following cortical injections of tritiated proline and/or horseradish peroxidase. All four structures were shown to receive topographically organized projections from the striate cortex. The exposed surface of the striate cortex was found to be connected to the rostral part of the SC and the caudal part of the LG. Injections of the exposed striate cortex close to its rostral border resulted in label in adjoining parts of the Pl and Pi. The ventral half and dorsal half of the calcarine fissure were connected with the medial and lateral parts of the SC, the ventrolateral and dorsomedial portions of the Pl and Pi and the lateral and medial parts of the LG, respectively. Injections located at the lateral posterior extreme of the calcarine fissure resulted in label at the optic disc representation in the LG. The horseradish peroxidase material demonstrated that LG neurons in all laminae and interlaminar zones project to the striate cortex.Abbreviations BIC brachium of the inferior colliculus - BSC brachium of the superior colliculus - C cerebellum - CG central grey - i interlaminar zone(s) of the dorsal lateral geniculate nucleus - IC inferior colliculus - ICc central nucleus of the inferior colliculus - LG dorsal lateral geniculate nucleus - m magnocellular layer(s) of the dorsal lateral geniculate nucleus - MG medial geniculate body - p parvocellular layer(s) of the dorsal lateral geniculate nucleus - P pulvinar complex - Pi inferior pulvinar - PG pregeniculate nucleus - Pl lateral pulvinar - Pm medial pulvinar - s superficial layer(s) of the dorsal lateral geniculate nucleus - SC superior colliculus - sgs stratum griseum superficiale of the superior colliculus - R reticular nucleus of the thalamus - VP ventroposterior group - 17 Area 17 Supported by NEI Grants EY-07007 (J. Graham) and EY-02686 (J.H. Kaas)     

Time of origin of neurons of the rat inferior colliculus and the relations between cytogenesis and tonotopic order in the auditory pathway

Summary Groups of pregnant rats were injected with two successive daily doses of ³H-thymidine from gestational day 12 and 13 (E12+13) until the day before parturition (E21+22) in order to label in their embryos the proliferating precursors of neurons. At 60 days of age the proportion of neurons generated (or no longer labelled) on specific embryonic days was determined quantitatively in six vertical strips of the inferior colliculus. It was established that the neurons of the inferior colliculus are produced between days E14 and the perinatal period in an orderly sequence: the earliest generated cells are situated rostrally, laterally and ventrally in the principal nucleus, the latest generated cells are situated caudally, medially and dorsally in the pericentral nucleus. This cytogenetic gradient suggested that the cells are produced dorsally in the caudal recess of the embryonic aqueduct and are deployed in an outsidein pattern.This study has brought to a conclusion our datings of neuron production in the central auditory pathway of the rat. The results revealed that in those structures in which a cytogenetic gradient could be recognized, the orientation of this gradient and the regional tonotopic order (demonstrated mostly in species other than the rat) tended to be aligned. Moreover, with the exception of the medial trapezoid nucleus and the dorsal nucleus of the lateral lemniscus (which receive contralateral input from the cochlear nuclei), sites with early-produced neurons correlated with units responding preferentially to high frequency tones and vice versa. This suggested that the orderly production of neurons within different components of the auditory system is a factor in their subsequent topographic organization. A comparison of the temporal order of neuron production in different components of the auditory pathway suggested that the establishment of orderly topographic relations between some of the structures (e.g., the medial geniculate body and the primary auditory cortex) takes place before this spatial relationship could be specified as a cochleotopic order.Abbreviations ab cochlear nerve, ascending branch - Ai aqueduct, inferior collicular recess - AI primary auditory cortex - bi brachium of inferior colliculus - c caudal - CE cerebellum - CI central nucleus, inferior colliculus - CNa anteroventral cochlear nucleus - CNd dorsal cochlear nucleus - CNp posteroventral cochlear nucleus - d dorsal - db cochlear nerve, descending branch - DI diencephalon - ds dorsal acoustic stria (stria of Monakow) - DM dorsomedial nucleus, inferior colliculus - EX external nucleus, inferior colliculus - IC inferior colliculus - is intermediate acoustic stria (stria of Held) - l lateral - LD dorsal nucleus of lateral lemniscus - ll lateral lemniscus - LV ventral nucleus of lateral lemniscus - m medial - ME medulla - MG medial geniculate body - MS mesencephalon - PC pericentral nucleus, inferior colliculus - PR principal nucleus, inferior colliculus - py pyramidal cells, dorsal cochlear nucleus - r rostral - SOl lateral superior olivary nucleus - SOm medial superior olivary nucleus - TRl lateral trapezoid nucleus - TRm medial trapezoid nucleus - v ventral - VL lateral ventricle - vs ventral acoustic stria (trapezoid body) - V3 third ventricle - VIIIn cochlear nerve     

Early blindness results in abnormal corticocortical and thalamocortical connections

Studies in congenitally blind and bilaterally enucleated individuals show that an early loss of sensory driven activity can lead to massive functional reorganization. However, the anatomical substrate for this functional reorganization is unknown. In the present study, we examined patterns of corticocortical and thalamocortical connections in adult opossums that had been bilaterally enucleated neonatally, prior to the formation of retinogeniculate and geniculocortical connections. We show that in addition to normal thalamocortical projection patterns from visual nuclei, enucleated animals also receive input from nuclei associated with the somatosensory (ventral posterior nucleus, VP), auditory (medial geniculate nucleus, MGN), motor (ventrolateral nucleus, VL), and limbic/hippocampal systems (anterior dorsal nucleus, AD; and anterior ventral nucleus, AV). Likewise, in addition to normal corticocortical projections to area 17, bilaterally enucleated opossums also receive input from auditory, somatosensory, and multimodal cortex. These aberrant patterns of thalamocortical and corticocortical connections can account for alterations in functional organization observed in the visual cortex of bilateral enucleated animals, and indicate that factors extrinsic to the cortex play a large role in cortical field development and evolution. On the other hand, the maintenance of normal patterns of connections in the absence of visual input suggests that there are formidable constraints imposed on the developing cortex that highly restrict the types of evolutionary change possible.     

Effect of acute and chronic bilateral visual deafferentation on c-Fos immunoreactivity in the visual system of adult rats

In our study we examined acute and chronic changes in c-Fos expression patterns in the visual system of the rat after complete visual deafferentation. In 20 male Lewis rats, the retro-bulbar part of the optic nerve was sectioned bilaterally. Ten animals underwent c-Fos immunohistochemistry after 3 days and 10 animals after 3 weeks examining time-dependent changes. The control group consisted of 10 animals, which did not undergo any surgical manipulation. c-Fos expression in the rat visual system experienced significant changes after acute and chronic bilateral complete visual deafferentation. Acute decrease in c-Fos level was observed in the ventral lateral geniculate nucleus, intergeniculate leaflet, superficial gray layer of the superior colliculus and layers IV and V of the primary visual cortex. After chronic deafferentation, c-Fos expression was also found to be decreased in the optic and deep layers of the superior colliculus and layer VI of the primary visual cortex. No change in c-Fos expression was observed in the dorsal lateral geniculate nucleus and layers I, II and III of the primary visual cortex. This work shows that secondary complete blindness does not lead to uniform decrease in c-Fos levels in all subcortical and cortical brain regions related to vision. These findings provide important information concerning expression of the immediate-early gene product c-Fos in secondary blind rodent models. It may further serve as a relevant baseline finding when electrical stimulation of the visual system is performed, aiding the assessment of visual neuroprosthesis using c-Fos as a functional mapping tool when evaluating different stimulus parameters in blind rodent models.     

Direct innervation of identified tectothalamic neurons in the inferior colliculus by axons from the cochlear nucleus.

The present study sought to identify tectothalamic neurons in the rat inferior colliculus that receive their innervation directly from the cochlear nuclei and to identify the axons that provide the innervation. A direct projection would bypass the binaural interactions of the superior olivary complex and provide the quickest route to the neocortex. Axons, primarily from the dorsal cochlear nucleus, were labeled with anterograde transport of dextran and terminated in the central nucleus of the inferior colliculus in a laminar pattern. Most labeled axons were thin and simply branched. Other axons were thicker, gnarly, less frequently observed and probably originated from the ventral cochlear nucleus. None had concentrated endbulbs or a nest of endings. Both types of axons terminated primarily in the central nucleus and layer 3 of the external cortex. This pattern suggests that the combination of these subdivisions in the rat are equivalent to the central nucleus as defined in other species. Tectothalamic neurons in the inferior colliculus in the same animals were identified by retrograde transport from the medial geniculate body and intracellular injection of Lucifer Yellow. A number of different cell types act as tectothalamic neurons and receive contacts from cochlear nucleus axons. These include flat cells (disc-shaped), less-flat cells and stellate cells. Two innervation patterns were seen: a combination of axosomatic and axodendritic contacts, and predominantly axodendritic contacts. Both patterns were seen in the central nucleus, but axosomatic contacts were seen less often in the other subdivisions. This is the first study to show direct connections between cochlear nuclear axons and identified tectothalamic neurons. The layers of axons from cochlear nuclei may provide convergent inputs to neurons in the inferior colliculus rather than the heavy inputs from single axons typical of lower auditory nuclei. Excitatory synapses made by axons from the cochlear nuclei on tectothalamic neurons may provide a substrate for rapid transmission of monaural information to the medial geniculate body.     

The auditory brainstem nuclei and some of their projections to the inferior colliculus in the North American opossum

Afferent projections to the inferior colliculus in the North American opossum have been examined using the retrograde transport of horseradish peroxidase. Projections to primarily the contralateral inferior colliculus arise in the dorsal and ventral cochlear nuclei, the auditory nerve nucleus and the spinal trigeminal nucleus pars caudalis, while ipsilateral projections arise in the superior paraolivary nucleus, the ventral nucleus of the trapezoid body, the ventral nucleus of the lateral lemniscus, the paralemniscal nucleus, the deep layer of the superior colliculus and the parabrachial nucleus. Bilateral projections to the inferior colliculus originate within the dorsal column nuclei, the nucleus reticularis gigantocellularis pars ventralis, the lateral and medial superior olivary nuclei, the dorsal nucleus of the lateral lemniscus and the auditory cortex. Nissl, fiber and Golgi-stained preparations were used to study the neuronal organization of those auditory nuclei with projections to the inferior colliculus. Anterograde axonal degeneration and transport techniques revealed that the inferior colliculus is innervated differentially by the dorsal and ventral cochlear nucleus, the superior olivary complex and the auditory neocortex. Axons from the contralateral dorsal cochlear nucleus and the ipsilateral superior olivary complex innervate both the central nucleus and external cortex, whereas those from ventral cochlear nucleus and contralateral, superior olivary complex project to only the central nucleus. Projections from auditory cortex form the complement of those from the cochlear nuclei and superior olivary complex, that is, they terminate in a thin band overlying the dorsal cortex and the superficial layer of external cortex.

Our results have been compared with those obtained from eutherian mammals and it is clear that there are striking similarities in neuronal organization and connectivity. Since the opossum is born 12 days after conception and has an extended development in an external pouch, it may be suited for developmental studies of the mammalian auditory connections and the behaviors dependent of them.     

Faster scaling of visual neurons in cortical areas relative to subcortical structures in non-human primate brains

Cortical expansion, both in absolute terms and in relation to subcortical structures, is considered a major trend in mammalian brain evolution with important functional implications, given that cortical computations should add complexity and flexibility to information processing. Here, we investigate the numbers of neurons that compose 4 structures in the visual pathway across 11 non-human primate species to determine the scaling relationships that apply to these structures and among them. We find that primary visual cortex, area V1, as well as the superior colliculus (SC) and lateral geniculate nucleus scale in mass faster than they gain neurons. Areas V1 and MT gain neurons proportionately to the entire cerebral cortex, and represent fairly constant proportions of all cortical neurons (36 and 3 %, respectively), while V1 gains neurons much faster than both subcortical structures examined. Larger primate brains therefore have increased ratios of cortical to subcortical neurons involved in processing visual information, as observed in the auditory pathway, but have a constant proportion of cortical neurons dedicated to the primary visual representation, and a fairly constant ratio of about 45 times more neurons in primary visual than in primary auditory cortical areas.     

Effects of stimulation of the primary auditory cortex upon colliculogeniculate neurons in the inferior colliculus of the cat

Electrical stimulation of the primary auditory cortex (AI) of the cat was found to evoke EPSPs, IPSPs or EPSP-IPSP sequences in colliculogeniculate (CG) neurons in the inferior colliculus (IC) which responded antidromically to stimulation of the medial geniculate nucleus. The CG neurons responding to the AI stimulation with short-latency EPSPs (1.0-1.4 msec) were located in the dorsomedial portion of the central nucleus of the IC. On the other hand, latencies of IPSPs elicited in CG neurons by AI stimulation ranged from 2.0 to 4.5 msec.     

Aberrant retinal projections in congenitally deaf mice: how are phenotypic characteristics specified in development and evolution?

The contribution of sensory input to the formation of sensory system-specific (sensoritopic) connections of the thalamus and midbrain was investigated using mice lacking the Na+-K+-2Cl- cotransporter (NKCC1) or the plasma membrane Ca2+-ATPase isoform2 (PMCA2). Because these mice are congenitally deaf, the developing nervous system has no exposure to sensory-driven neural activity from the auditory system. Here we compared the retinofugal pathway in normal and congenitally deaf mice using intraocular injections of neuroanatomical tracers into each eye, and relating tracer patterns to identified thalamic nuclei and superior colliculus layers. We demonstrate that loss of such activity results in aberrant projections of the retina into nonvisual auditory structures such as the medial geniculate nucleus and the intermediate layers of the superior colliculus. These findings indicate that activity from peripheral sensory receptor arrays is necessary not only for the refinement of developing connections within a unimodal structure, but for the establishment of sensoritopic or sensory-specific connections of unimodal and multimodal structures. We hypothesize that specification of such connections may occur through the modulation of spatial expression patterns of molecules known to be involved in the development of topography of connections between brain structures, such as the ephrins, via activity-dependent, CRE-mediated gene expression.     

Multisensory processing via early cortical stages: Connections of the primary auditory cortical field with other sensory systems

It is still a popular view that primary sensory cortices are unimodal, but recent physiological studies have shown that under certain behavioral conditions primary sensory cortices can also be activated by multiple other modalities. Here, we investigate the anatomical substrate, which may underlie multisensory processes at the level of the primary auditory cortex (field AI), and which may, in turn, enable AI to influence other sensory systems. We approached this issue by means of the axonal transport of the sensitive bidirectional neuronal tracer fluorescein-labeled dextran which was injected into AI of Mongolian gerbils (Meriones unguiculatus). Of the total number of retrogradely labeled cell bodies (i.e. cells of origin of direct projections to AI) found in non-auditory sensory and multisensory brain areas, approximately 40% were in cortical areas and 60% in subcortical structures. Of the cell bodies in the cortical areas about 82% were located in multisensory cortex, viz., the dorsoposterior and ventroposterior, posterior parietal cortex, the claustrum, and the endopiriform nucleus, 10% were located in the primary somatosensory cortex (hindlimb and trunk region), and 8% in secondary visual cortex. The cortical regions with retrogradely labeled cells also contained anterogradely labeled axons and their terminations, i.e. they are also target areas of direct projections from AI. In addition, the primary olfactory cortex was identified as a target area of projections from AI. The laminar pattern of corticocortical connections suggests that AI receives primarily cortical feedback-type inputs and projects in a feedforward manner to its target areas. Of the labeled cell bodies in the subcortical structures, approximately 90% were located in multisensory thalamic, 4% in visual thalamic, and 6% in multisensory lower brainstem structures. At subcortical levels, we observed a similar correspondence of retrogradely labeled cells and anterogradely labeled axons and terminals in visual (posterior limitans thalamic nucleus) and multisensory thalamic nuclei (dorsal and medial division of the medial geniculate body, suprageniculate nucleus, posterior thalamic cell group, zona incerta), and in the multisensory nucleus of the brachium of the inferior colliculus. Retrograde, but not anterograde, labeling was found in the multisensory pontine reticular formation, particularly in the reticulotegmental nucleus of the pons. Conversely, anterograde, but no retrograde, labeling was found in the visual laterodorsal and lateroposterior thalamic nuclei, in the multisensory peripeduncular, posterior intralaminar, and reticular thalamic nuclei, as well as in the multisensory superior and pericentral inferior colliculi (including cuneiform and sagulum nucleus), pontine nuclei, and periaqueductal gray. Our study supports the notion that AI is not merely involved in the analysis of auditory stimulus properties but also in processing of other sensory and multisensory information. Since AI is directly connected to other primary sensory cortices (viz. the somatosensory and olfactory ones) multisensory information is probably also processed in these cortices. This suggests more generally, that primary sensory cortices may not be unimodal.     

Projections to the inferior colliculus from layer VI cells of auditory cortex

A large injection of a retrograde tracer into the inferior colliculus of guinea pigs labeled two bands of cells in the ipsilateral auditory cortex: a dense band of cells in layer V and a second band of cells in layer VI. On the contralateral side, labeled cells were restricted to layer V. The ipsilateral layer VI cells were distributed throughout temporal cortex, suggesting projections from multiple auditory areas. The layer VI cells included pyramidal cells as well as several varieties of non-pyramidal cells. Small tracer injections restricted to the dorsal cortex or external cortex of the inferior colliculus consistently labeled cells in layer VI. Injections restricted to the central nucleus of the inferior colliculus labeled layer VI cells only rarely. Overall, 10% of the cells in temporal cortex that project to the ipsilateral inferior colliculus were located in layer VI, suggesting that layer VI cells make a significant contribution to the corticocollicular pathway.     

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.     

Estrogen receptors in the central auditory system of male and female mice

The estrogen receptors in the central auditory system of male and female mice were characterized using immunocytochemical methods. Estrogen receptors α and β (ERα, ERβ) were localized predominantly in the ventral cochlear nucleus, nucleus of the trapezoid body, the lateral- and medio-ventral periolivary nuclei, the dorsal lateral lemniscus, and the inferior colliculus. The medial geniculate nucleus was negative for both ERα and ERβ whereas the auditory cortex was positive for ERα. The lateral superior olive, the ventral lateral lemniscus and the central nucleus of the inferior colliculus expressed only ERβ. The differential localization of ERα and ERβ may indicate distinct roles for these two receptors in auditory processing. No major differences in the pattern, number or intensity of receptor expression was found between male and female animals. The comprehensive anatomic map that is constructed for ERα and ERβ in the central auditory pathway will be a useful foundation to elucidate the complexity of estrogen actions in the auditory system.     

Cortical and subcortical connections of the pars compacta of the anterior pretectal nucleus in the rat.

The efferent and afferent connections of the dorsal part of the anterior pretectal nucleus, pars compacta (APc), were studied experimentally in the rat by using neurotracers. A restricted number of structures supply afferents to the anterior pretectal nucleus: the visual cortex (areas 17, 18 and 18a), ventral lateral geniculate nucleus and superficial layers of the superior colliculus. Additional afferents have been demonstrated originating from the Darkschewitsch nucleus, periaqueductal gray, zona incerta and anterior cingulate cortex. Efferent fibers are distributed to a sector of the deep mesencephalic nucleus just dorsolateral to the red nucleus, the basilar pontine gray, posterior and olivary pretectal nuclei, superficial layers of the superior colliculus, lateral posterior thalamic nucleus, ventral lateral geniculate nucleus and zona incerta. These anatomical observations indicate that the pars compacta of the anterior pretectal nucleus is closely related to visual centers, suggesting an involvement of this nucleus in visually mediated behavior.