Pathways from auditory cortex to the cochlear nucleus in guinea pigs.

The inferior colliculus (IC) and superior olivary complex (SOC) are important sources of descending pathways to the cochlear nucleus. The IC and SOC are also targets of direct projections from the auditory cortex but it is not known if cortical axons contact the cells that project to the cochlear nucleus. Multi-labeling techniques were used to address this question in guinea pigs. A fluorescent anterograde tracer was injected into temporal cortex to label corticofugal axons. Different fluorescent tracers were injected into one or both cochlear nuclei to label olivary and collicular cells. The brain was subsequently processed for fluorescence microscopy and the IC and SOC were examined for apparent contacts between cortical axons and retrogradely labeled cells. The results suggest that cortical axons contact cochlear nucleus-projecting cells in both IC and SOC. In both regions, contacts were more numerous on the side ipsilateral to the injected cortex. In the IC, the contacted cells projected ipsilaterally or contralaterally to the CN. In the SOC, the contacted cells projected ipsilaterally, contralaterally or bilaterally to the CN. We conclude that auditory cortex is in a position to modulate descending pathways from both the IC and SOC to the cochlear nucleus.

Cells in auditory cortex that project to the cochlear nucleus in guinea pigs.

Fluorescent retrograde tracers were used to identify the cells in auditory cortex that project directly to the cochlear nucleus (CN). Following injection of a tracer into the CN, cells were labeled bilaterally in primary auditory cortex and the dorsocaudal auditory field as well as several surrounding fields. On both sides, the cells were limited to layer V. The size of labeled cell bodies varied considerably, suggesting that different cell types may project to the CN. Cells ranging from small to medium in size were present bilaterally, whereas the largest cells were labeled only ipsilaterally. In optimal cases, the extent of dendritic labeling was sufficient to identify the morphologic class. Many cells had an apical dendrite that could be traced to a terminal tuft in layer I. Such "tufted" pyramidal cells were identified both ipsilateral and contralateral to the injected CN. The results suggest that the direct pathway from auditory cortex to the cochlear nucleus is substantial and is likely to play a role in modulating the way the cochlear nucleus processes acoustic stimuli.

Projections from auditory cortex contact cells in the cochlear nucleus that project to the inferior colliculus.

Anterograde and retrograde tracing techniques were combined to determine whether auditory cortical axons contact cells in the cochlear nucleus that project to the inferior colliculus. FluoroRuby or fluorescein dextran was injected into auditory cortex to label cortical axons by anterograde transport. Different fluorescent tracers (Fast Blue, FluoroGold, FluoroRuby or fluorescein dextran) were injected into one or both inferior colliculi to label cells in the cochlear nucleus. After 12-15 days, the brain was processed for fluorescence microscopy and the cochlear nuclei were examined for apparent contacts between cortical axons and retrogradely labeled cochlear nucleus cells. The results suggest that axons from the ipsilateral or contralateral cortex contact fusiform and giant cells in the dorsal cochlear nucleus and multipolar cells in the ventral cochlear nucleus that project directly to the inferior colliculus. The contacts occur on cell bodies and dendrites. The target cells in the cochlear nucleus include cells that project ipsilaterally, contralaterally or bilaterally to the inferior colliculus. The results suggest that auditory cortex is in a position to exert direct effects on the monaural pathways that ascend from the cochlear nucleus.

Unilateral and bilateral projections from cortical cells to the inferior colliculus in guinea pigs.

Auditory cortex projects directly and bilaterally to the inferior colliculus (IC). We used multiple fluorescent retrograde tracers to determine whether individual cortical cells project to both the left and right IC. Injection of different tracers into each IC labeled many cells in a sheet that extended throughout much of temporal cortex in both hemispheres. Most cells contained a single tracer, with the majority of these labeled from the ipsilateral IC. Numerous double-labeled cells were observed throughout the same areas of temporal cortex. The double-labeled cells form a small percentage of the cortical cells that project to the ipsilateral IC (6.1% on average) and a much larger percentage of the cells that project to the contralateral IC (46.4% on average). Unilaterally projecting cells are well positioned to have effects limited to one IC, whereas bilaterally projecting cells are likely to have a broader influence and may coordinate activity on the two sides of the midbrain.

Auditory cortical projections to the cochlear nucleus in guinea pigs.

We used anterograde tracing techniques to examine projections from auditory cortex to the cochlear nucleus in guinea pigs. Following injection of dextrans into the temporal cortex, labeled axons were present bilaterally in the cochlear nucleus. The distribution of boutons within the cochlear nucleus was similar on the two sides. The majority of boutons was usually located on the ipsilateral side. Most of the boutons were located in the granule cell areas, where many small boutons and a few larger, mossy-type endings were labeled. Additional small, labeled boutons were found in all layers of the dorsal cochlear nucleus, with the majority located in the fusiform cell layer. Labeled boutons were also present in the ventral cochlear nucleus, where they were located in the small cell cap as well as magnocellular parts of both posteroventral and anteroventral cochlear nucleus. Similar results were obtained with injections restricted to primary auditory cortex or to the dorsocaudal auditory field. The results illustrate direct cortical projections to the cochlear nucleus that are likely to modulate the activity in a number of ascending auditory pathways.

Separate projections from the inferior colliculus to the cochlear nucleus and thalamus in guinea pigs.

We used multiple-labeling techniques with retrograde fluorescent tracers to determine whether individual cells in the inferior colliculus project to the medial geniculate body (MG) and the cochlear nucleus (CN) in guinea pigs. Four possible projection patterns were examined: (1) to ipsilateral MG and ipsilateral CN; (2) to ipsilateral MG and contralateral CN; (3) to contralateral MG and ipsilateral CN; and, (4) to contralateral MG and contralateral CN. Following injections of different tracers into two or more sites, no inferior collicular cells were double-labeled from the two contralateral targets and only a few cells were double-labeled from each of the other pairs of targets. The double-labeled cells always totaled < 1% of the single-labeled populations. We conclude that collateral projections from the inferior colliculus to the MG and CN are virtually non-existent. Therefore, the ascending and descending projections to these targets arise from different cells. These cells could potentially receive different inputs and send different information to higher or lower centers of the auditory pathway.

Projections from the auditory cortex to the superior olivary complex in guinea pigs.

We used anterograde tracing techniques to characterize projections from auditory cortex to the superior olivary complex (SOC) in guinea pigs. Large injections of fluorescent or biotinylated dextrans into the temporal cortex labeled many axons in the SOC. Labeled boutons were most numerous in the ventral nucleus of the trapezoid body, with additional boutons in all other olivary nuclei. The distribution of boutons was similar in the ipsilateral and contralateral SOC; however, the contralateral SOC had markedly fewer axons and boutons. Similar patterns of labeling were also observed following injections confined to primary auditory cortex or the dorsocaudal auditory field. Cortical axons in many of the SOC nuclei share numerous morphological features, suggesting that individual axons may innervate multiple nuclei and have widespread effects. In addition, some nuclei contain axons with branching or termination patterns unique to that nucleus; these axons may represent focused projections with effects limited to individual SOC nuclei. Given the many projections of SOC nuclei, cortico-olivary projections are in a position to modify the activity of many brainstem auditory circuits.

GABAergic circuitry in the dorsal division of the cat medial geniculate nucleus.

Gamma-aminobutyric acid (GABA) is the main inhibitory neurotransmitter of the thalamus. We used postembedding immunocytochemistry to examine the synaptic organization of GABA-positive profiles in the dorsal superficial subdivision of the cat medial geniculate nucleus (MGN). Three groups of GABA-positive profiles participate in synapses: axon terminals, dendrites, and presynaptic dendrites. The presynaptic GABA-positive terminals target mainly GABA-negative dendrites. The GABA-positive postsynaptic profiles receive input primarily from GABA-negative axons. The results indicate that the synaptic organization of GABA-positive profiles in the dorsal superficial subdivision of the MGN nucleus is very similar to that in other thalamic nuclei.