Corticostriatal projections from layer V cells in rat are collaterals of long-range corticofugal axons

Corticostriatal projections arising from the infragranular layers of the motor and second somatosensory cortices were studied in rats after labeling small pools of neurons with biocytin. Camera lucida reconstruction of 263 fibers arising from laminae V and VI revealed that all corticostriatal projections derive from collaterals of lamina V cells whose main axons descend into the cerebral peduncle. In contrast, lamina VI cells do not branch upon the striatum, but upon the thalamus. Together with the results obtained in previous tracing studies, the present data raise the possibility that no neuron is exclusively corticostriatal. We therefore propose that all corticostriatal projections are collaterals given off by the axons of two types of neurons: layer V cells whose main axon project to the brainstem and/or spinal cord, and layer III cells that project to the contralateral hemisphere.     

Time Course of Inhibition Induced by a Putative Saccadic Suppression Circuit in the Dorsal Lateral Geniculate Nucleus of the Rabbit

Psychological studies have revealed that a visual suppression occurs during the saccadic eye movements to maintain the stable visual image. This visual suppression is named saccadic suppression. A typical saccadic suppression precedes the saccadic eye movements by 30–60 ms, lasts 120–180 ms, and is followed by a 100–150-ms facilitation. Recently, we have revealed an inhibitory circuit connecting the deep layers of the superior colliculus (SC) to the dorsal lateral geniculate nucleus (LGN), via the central lateral nucleus in the thalamus (CL) and thalamic reticular nucleus (TRN). We speculated that this inhibitory circuit might mediate saccadic suppression in the rabbit. In the present study, we used intracellular recording technique to further examine the synaptic and intrinsic responses of CL cells, TRN cells, and LGN cells to the activation of this inhibitory circuit. We found that the stimulation of the deeper layers of the SC induced a fast excitation postsynaptic potential (EPSP) in CL cells, followed by a robust EPSP in TRN cells and a prolonged inhibitory postsynaptic potential (IPSP) in LGN cells. The EPSP in TRN cells was always followed by a small inhibitory postsynaptic potential (IPSP). The IPSP in LGN cells lasted about 133 ± 27 ms. Sometimes, a rebound bursting occurred after the IPSP in LGN cells. We also examined whether activation of this inhibitory circuit could suppress the retino-geniculo-cortical pathway. We found that the SC stimulation always suppressed the evoked potential in the visual cortex induced by the stimulation of the optic chiasm. Our results of the inhibitory circuit can induce an inhibition in the LGN and a suppression on the retino-geniculo-cortical pathway. The time courses of the inhibition and suppression were compatible with that of saccadic suppression revealed by psychological and physiological studies. These results support the idea that the inhibitory circuit of SC (deeper layers)-CL-TRN-LGN may mediate the saccadic suppression in the rabbit LGN. Copyright © 1996 Elsevier Science Inc.     

Correlation between different types of retinal ganglion cells and their projection pattern in the albino rat

Injecting Fluoro-Gold (FG) and Evans-Blue (EB) into the right dLGN and SC in the adult albino rat, ipsilaterally projecting double-labeled retinal ganglion cells were mainly seen in the ventrotemporal crescent. They were mainly large sized cells. The ipsilaterally projecting double-labeled cells tended to have larger somata than the single- and double-labeled cells projecting to the contralateral superior colliculus and/or dorsal nucleus of the lateral geniculate body.     

Mode of termination of afferents from the thalamic reticular nucleus in the dorsal lateral geniculate nucleus of the rat

The terminals of axons projecting to the dorsal lateral geniculate nucleus from the thalamic reticular nucleus were identified by electron microscopy 8–24 h after placing small lesions in the ipsilateral reticular nucleus. The terminals contained flattened synaptic vesicles and made Gray type II axo-dendritic synaptic contacts with geniculate neurons. Their identification as F-axons accords well with physiological evidence for a powerful monosynaptic inhibitory input to geniculocortical projection cells from reticular nucleus neurons.     

Excitotoxic lesions of the rostral thalamic reticular nucleus do not affect the performance of spatial learning and memory tasks in the rat

Rats with cytotoxic lesions of the rostral pole of the thalamic reticular nucleus were compared with surgical control animals on a series of spatial learning and memory tests. While evidence was found for an initial, transient impairment on forced-choice alternation in a T-maze, this rapidly disappeared, and overall performance was unaffected. Subsequent experiments found no evidence that lesions of the rostral reticular nucleus affected the acquisition or performance of tests in the radial arm maze and the Morris water maze. Thus, it appears that the rostral pole of the thalamic reticular nucleus often does not play a necessary role in the performance of tests of spatial learning and memory, in spite of its interconnections with other regions that are required for normal spatial memory.     

The conduction velocity and central projections of retinofugal fibers in the rabbit

In anesthetized and paralyzed rabbits, action potentials were elicited antidromically following electrical stimulation of optic tract terminals at the geniculate level (LGN). The conduction velocity spectrum extended from 7 m.s⁻¹ to 34 m.s⁻¹. The distribution of conduction velocities indicated four major modes at 10, 18, 22 and 26 m.s⁻¹. Antidromic compound action potentials exhibited good correlation between the conduction latency and the major modes of the distribution histogram. These results suggest that the rabbit optic tract is composed of four classes of fibers varying in their conduction velocities. The central projections of retinofugal axons were studied with electrical stimulations of the Superior Colliculus (CS) and the LGN while recording from the same optic tract fiber. Antidromic spikes could be elicited from all conduction velocity groups and 71% of axons responded to both sites of stimulation. This finding indicates that most retinofugal fibers branch to and innervate both CS and LGN. Further, there is a tendency for fast-conducting axons to have their receptive field located eccentrically relative to the optic axis of the eye.     

Latency distribution from orthodromic stimulation at the optic nerve, in the Lateral Geniculate Nucleus and Superior Colliculus of rabbits

Latency distribution of responses to optic nerve stimulation has been compared between the Lateral Geniculate Nucleus (L.G.N.) and Superior Colliculus (S.C.), in rabbits. Histograms revealed that in both structures the latency distributions are very similar. Results suggest that all groups of retinofugal fibers project to both sites, although longer latencies are more frequently met at the collicular level. In addition evidence is presented that in rabbit's optic nerve there are at least three populations of axons with different conduction velocities.     

Development of the projections from the dorsal lateral geniculate nucleus to the lateral suprasylvian visual area of cortex in the cat.

In the study reported in the preceding paper, we used retrograde labeling methods to show that the enhanced projection from the thalamus to the posteromedial lateral suprasylvian (PMLS) visual area of cortex that is present in adult cats following neonatal visual cortex damage arises at least partly from surviving neurons in the dorsal lateral geniculate nucleus (LGN). In the C layers of the LGN, many more cells than normal are retrogradely labeled by horseradish peroxidase (HRP) injected into PMLS cortex ipsilateral to a visual cortex lesion. In addition, retrogradely labeled cells are found in the A layers, which normally have no projection to PMLS cortex in adult cats. The purpose of the present study was to investigate the mechanisms of this enhanced projection by examining the normal development of projections from the thalamus, especially the LGN, to PMLS cortex. Injections of HRP were made into PMLS cortex on the day of birth or at 1, 2, 4, or 8 weeks of age. Retrogradely labeled neurons were present in the lateral posterior nucleus, posterior nucleus of Rioch, pulvinar, and medial interlaminar nucleus, as well as in the LGN, at all ages studied. Within the LGN of the youngest kittens, a small number of retrogradely labeled cells was present in the interlaminar zones and among the cells in the A layers that border these zones. Such labeled cells were virtually absent by 8 weeks of age, and they are not found in normal adult cats. Sparse retrograde labeling of C-layer neurons also was present in newborn kittens.(ABSTRACT TRUNCATED AT 250 WORDS)     

Physiological properties of the output neurons in the deep layers of the superior colliculus of the rabbit

Using antidromic and orthodromic stimulation techniques, we studied physiological properties of the output neurons in the deep layers of the superior colliculus (SC) of 34 New Zealand rabbits. SC cells antidromicaly activated from the contralateral predorsal bundle (PDB) could also be activated by stimulation of the contralateral SC and ipsilateral central lateral nucleus of the thalamus (CL). The majority of these output neurons responded predominantly to the stimulation of the optic nerve, and only a small proportion of the output neurons were responsive to the stimulation of somatosensory and auditory (and/or vestibular) nerves. These results suggest that the orienting reflex might be elicited mainly by visual afferents in the rabbit The output SC neurons were subject to a 70 ms inhibition after antidromic stimulation of the PDB and a 40 ms inhibition after transsynaptic (orthodromic) stimulation of the optic chiasm (OX), indicating that the output neurons in the deep layers of the SC might be subject to at least two inhibitory circuits. These results are discussed in the context of a putative saccadic suppression circuitry model.     

Localization of two high-threshold potassium channel subunits in the rat central auditory system

The firing pattern of auditory neurons is determined in part by the type of voltage-sensitive potassium channels expressed. The expression patterns for two high-threshold potassium channels, Kv3.1 and Kv3.3, that differ in inactivation properties were examined in the rat auditory system. The positive activation voltage and rapid deactivation kinetics of these channels provide rapid repolarization of action potentials with little effect on action potential threshold. In situ hybridization experiments showed that Kv3.3 mRNA was highly expressed in most auditory neurons in the rat brainstem, whereas Kv3.1 was expressed in a more limited population of auditory neurons. Notably, Kv3.1 mRNA was not expressed in neurons of the medial and lateral superior olive and a subpopulation of neurons in the ventral nucleus of the lateral lemniscus. These results suggest that Kv3.3 channels may be the dominant Kv3 subfamily member expressed in brainstem auditory neurons and that, in some auditory neurons, Kv3.1 and Kv3.3 may coassemble to form functional channels. The localization of Kv3.1 protein was examined immunohistochemically. The distribution of stained somata and neuropil varied across auditory nuclei and correlated with the distribution of Kv3.1 mRNA-expressing neurons and their terminal arborizations, respectively. The intensity of Kv3.1 immunoreactivity varied across the tonotopic map in the medial nucleus of the trapezoid body with neurons responding best to high-frequency tones most intensely labeled. Thus, auditory neurons may vary the types and amount of K(+) channel expression in response to synaptic input to subtly tune their firing properties.     

Calcitonin gene-related peptide (CGRP) immunoreactive projections from the thalamus to the striatum and amygdala in the rat

The organization of calcitonin gene-related peptide-like immunoreactive (CGRPir) innervation of the amygdala and caudate-putamen in the rat was examined by using immunohistochemistry for CGRP combined with retrograde transport of the fluorescent dye fluoro-gold, as well as anterograde transport of Phaseoleus vulgaris leucoagglutinin (PHA-L). The lateral part of the central nucleus of the amygdala and the amygdalostriatal transition zone was densely innervated by CGRPir terminals at all anterior-posterior levels. More caudally, the lateral part of the caudate-putamen also had large numbers of CGRPir terminals. Injections of fluoro-gold into the amygdala and amygdalostriatal transition area followed by immunohistochemistry for CGRP revealed double-labeled neurons in the subparafascicular, lateral subparafascicular, and posterior intralaminar nuclei of the thalamus and peripeduncular nucleus. Injections into the caudate-putamen demonstrated double-labeled neurons in the more lateral parts of this same nuclear complex. PHA-L injections into the posterior thalamic nuclei from which the CGRPir projections arise confirmed the medial-to-lateral organization of the projections to the amygdala and striatum. The subparafascicular nucleus and the rostral portion of the lateral subparafascicular nucleus primarily projected to the medial amygdala and the amygdalostriatal transition area, while the more lateral cell groups, including the caudal part of the lateral parafascicular, posterior intralaminar, and peripeduncular nuclei projected to the lateral amygdala and the caudate-putamen. These CGRPir projections may be involved in mediating conditioned autonomic and behavioral responses to acoustic stimuli or somatosensory stimuli.     

Serotonin-mediated inhibition from dorsal raphe nucleus of neurons in dorsal lateral geniculate and thalamic reticular nuclei

Electrophysiological studies using rats anesthetized with chloral hydrate were performed to determine whether or not serotonin originating in the dorsal raphe nucleus (DR) acts as an inhibitory transmitter or neuromodulator on neurons of the dorsal lateral geniculate nucleus (LGN) and neurons located in the thalamic reticular nucleus (TRN) immediately rostral to the dorsal LGN. In the LGN, conditioning stimuli applied to the DR preceding test stimulus to the optic tract and visual cortex inhibited orthodromic and antidromic spikes in about one-third of the relay neurons and in more than half of the intrageniculate interneurons. Conditioning stimulation of the DR also produced an inhibition of the spikes elicited by stimulation of the optic tract and visual cortex of at least three-quarters of the TRN neurons. Iontophoretic application of serotonin (25 nA) inhibited the orthodromic spikes of the LGN relay neuron and TRN neuron. A close correlation was observed between the effects of DR conditioning stimulation and iontophoretic serotonin in the same neurons. The inhibition with DR conditioning stimulation and iontophoretically applied serotonin was antagonized during iontophoretic application of methysergide (15-40 nA), a serotonin antagonist. These results strongly suggest that serotonin derived from the DR acts on the LGN and TRN neurons as an inhibitory transmitter or neuromodulator to inhibit transmission in these nuclei.     

Effect of optic nerve transection onN-acetylaspartylglutamate immunoreactivity in the primary and accessory optic projection systems in the rat

Evidence has been presented in recent years that support the hypothesis thatN-acetylaspartylglutamate (NAAG) may be involved in synaptic transmission in the optic tract of mammals. Using a modified fixation protocol, we have determined the detailed distribution of NAAG immunoreactivity (NAAG-IR) in retinal ganglion cells and optic projections of the rat. Following optic nerve transection, dramatic losses of NAAG-IR were observed in the neuropil of all retinal target zones including the lateral geniculate nucleus, superior colliculus, nucleus of the optic tract, the dorsal and medial terminal nuclei and suprachiasmatic nucleus. Brain regions were microdissected and NAAG levels measured by a radioimmunoassay (RIA) (IC₅₀:NAAG= 2.5nM,NAA= 100 μM;smallest detectable amount= 1–2pg/assay). decreases (50–60%) in NAAG levels were detected in the lateral geniculate, superior colliculus and suprachiasmatic nucleus. Moderate losses (25–45%) were noted in the pretectal nucleus and the nucleus of the optic tract. Smaller changes (15–20%) were detected in the paraventricular nucleus and the pretectal area. These results are consistent with a synaptic communication role for NAAG in the visual system.     

Topographic organization of projections from the thalamic reticular nucleus to the anterior thalamic nuclei in the rat

We have investigated connections between the thalamic reticular nucleus (TRN) and the anterior thalamic nuclei (ATN) in the rat, following injections of horseradish peroxidase (HRP) into subnuclei of the ATN and different regions of the rostral TRN. Three nonoverlapping groups of neurons in the dorsal part of the ipsilateral rostral TRN project to, and receive reciprocal projections from, specific subnuclei of the ATN. A vertical sheet of neurons in the most dorsal part of the rostral TRN projects to the dorsal half of the posterior subdivision of the anteroventral thalamic nucleus (AVp), the dorsal region of the medial subdivision of the anteroventral thalamic nucleus (AVm), and the dorsolateral part of the rostral anterodorsal thalamic nucleus (AD). Immediately ventral to this part of TRN, but still within its dorsal portion, are a lateral cluster of neurons and a medially located vertical sheet of neurons. The lateral cluster projects to the ventral part of AVp and to the dorsomedial part of rostral AD. The medial sheet projects to the ventral part of AVm, the ventral part of rostral AD, and to the caudal portions of both AV and AD. There appears to be no input to the anteromedial thalamic nucleus (AM) from the TRN. These findings shed new light on the anatomy of the rostral TRN, the ATN, and the connections between the two, and are relevant to emerging hypotheses about the functional organization of the TRN and reticulo-thalamic projections.     

Organization of projections from the medial agranular cortex to the superior colliculus in the rat: a study using anterograde and retrograde tracing methods

The organization of corticotectal projections from the medial agranular cortex (AGm), which has been considered to contain rat's frontal eye field, was examined using anterograde and retrograde tracing techniques. When biotinylated dextranamine (BDA) injections were made into the rostral part of the AGm, small numbers of BDA-labeled axons were found in the rostral two-thirds of the superior colliculus (SC) while some labeled axons were seen in the caudal one-third of the SC. These labeled axons were distributed mainly in the lateral part of the stratum griseum intermediale. On the other hand, after BDA injections into the caudal part of the AGm, moderate to dense plexuses of labeled axons were found in the rostral two-thirds of the SC while some labeled axons were seen in the caudal one-third of the SC. These labeled axons were distributed in the ventromedial and dorsolateral marginal zones of the stratum griseum intermediale as well as in the stratum griseum profundum. The corticotectal projections were largely uncrossed. After combined injections of BDA into the caudal part of the AGm on one side and cholera toxin B subunit (CTb) into the paramedian pontine reticular formation on the opposite side or into the interstitial nucleus of Cajal on the same side, the overlapping distributions of BDA-labeled axons and CTb-labeled neurons were found in the ventromedial marginal zone of the stratum griseum intermediale ipsilateral to the site of BDA injection. These results suggest that the caudal part of the AGm plays a more significant role in the oculomotor function than does the rostral part of the AGm.     

Ipsilateral retinal projections and laminations of the dorsal lateral geniculate nucleus in the eastern chipmunk (Tamias sibiricus asiaticus)

The ipsilateral retinal projections and laminations of the dorsal geniculate nucleus (LGNd) were studied in the eastern chipmunk (Tamias sibiricus asiaticus). From cyto-, myelo- and chemoarchitectures the LGNd was divided into the ventromedial and dorsolateral parts. Anterograde axonal transport of wheat germ agglutinated horseradish peroxidase (WGA-HRP) and Fink-Heimer staining after unilateral eye removal both indicated that the ipsilateral projection terminates in lamina 2 of the ventromedial part and in lamina 3b of the dorsalateral part. The remaining laminas, lamina 1 of the ventromedial part and laminas 3a and 3c of the dorsolateral part, receive afferents from the contralateral retina. It was suggested that laminas 1 and 2 of the chipmunk's LGNd correspond to laminas A and A1 of the cat's LGNd, and the lamina 3 complex to its lamina C complex.     

Demonstration of axon collateral projections from the substantia nigra pars reticulata to the superior colliculus and the parvicellular reticular formation in the rat

It was revealed in the rat that single neurons in the substantia nigra pars reticulata (SNr) innervated both the superior colliculus (SC) and the parvicellular reticular formation (RFp) in the pons and medulla oblongata by way of axon collaterals. After injecting Fluoro-gold into the lateral part of the SC and Fluoro-ruby into the RFp on the same side, some SNr neurons were double-labeled with both tracers. They were localized in the dorsolateral part of the caudal half of the SNr ipsilateral to the injection sites.     

Cortical modulation of neuronal activity in the cat's lateral geniculate and perigeniculate nuclei

The cortico-thalamic influence on spontaneous and visually evoked activity of single cells in the dorsal lateral geniculate (LGN) and perigeniculate (PGN) nuclei were examined in unanesthetized cats with pretrigeminal brainstem transections by means of reversible cooling of cortical areas 17 and 18. The spatio-temporal characteristic of cells' RFs was tested with light spot randomly presented at different points along the receptive field axis. The cessation of cortical input decreased spontaneous activity of most of the LGN cells (64%; as compared to 36% with increased background firing). Similarly, their visually evoked responses were reduced (70% cells; compared to 24% with increased response) and extent of central excitatory domains diminished. In contrast, the majority of PGN neurons increased their spontaneous activity (62%; compared to 38% with decreased firing rate). Cortical cooling resulted also in a decrease of the ON and OFF central responses of most PGN cells (55%; as compared to 20% with increased responses). The described effects were more pronounced within the population of cells in X than in Y pathway. Although the removal of descending cortical excitation disturbed the balance of activity within the network of thalamic cells the gain of the geniculate relay was preserved. We conclude that the main role exerted by the cortico-thalamic pathway serves facilitation of the ascending retino-cortical flow of visual information at the level of lateral geniculate nucleus.