Central Lateral Nucleus Research Articles

Substance P innervation of the rat and cat thalamus. I. Distribution and relation to ascending spinal pathways.

An antiserum for substance P (SP) with minimal cross-reactivity for other tachykinins was employed to map the distribution of SP-positive nerve fibers and terminals in the thalamus of cats and rats with special emphasis on the innervation by these fibers of nuclei related to the somatosensory system. In both species SP innervation is predominantly along the midline, in medial and posterior thalamic regions, and sparser in sensory relays for specific modalities. Among the most densely innervated nuclei are the parafascicular, paraventricular, rhomboid, central medial and parts of mediodorsal, lateral posterior, and ventral lateral geniculate. SP innervation of somatosensory-related nuclei is also evident in central lateral nucleus, posterior complex (PO), and in ventroposterolateral (VPL) nucleus of both cats and rats. In VPL of cats SP fibers and terminals are present along its ventral and lateral border, a paralaminar area in which spinothalamic fibers have been shown to terminate and where neurons responsive to noxious stimuli have been reported. Also in rats the SP innervation of VPL is similar to that of spinothalamic tract fibers. The SP innervation of somatosensory thalamic nuclei may be supplied, at least in part, by spinothalamic afferent as suggested by the depletion of SP after anterolateral chordotomy but not after ablation of the dorsal column nuclei. The presence of SP-positive spinothalamic neurons in the spinal cord is reported in the following paper.

Peripheral and central components of alerting: Habituation of acoustic startle, orienting responses, and elicited waveforms.

Behavioral orienting (OR), the acoustic startle reflex (ASR), pontogeniculooccipital (PGO) waves in the lateral geniculate body, and midlatency auditory evoked responses (MLR) represent components of alerting. The habituation rate for each was examined to test the hypothesis that OR, ASR, and PGO waves have related underlying neural mechanisms and determine the similarity in responsiveness between elicited PGO waves (PGOE) and elicited waves in the thalamic central lateral nucleus (CLE), a site that yields MLR. PGOE and CLE waves did not habituate in amplitude after 120 tones; however, the pattern of responses for each waveform was different. OR and ASR significantly decreased amplitude across trials with OR exhibiting a faster, more pronounced decrement. Some separation exists between the peripheral (OR and ASR) and central (PGOE and CLE) components of alerting. PGO and CL waves may have common underlying neural mechanisms.

Sensory responses of intralaminar thalamic neurons activated by the superior colliculus.

The intralaminar thalamus of anesthetized rats was explored for neurons activated by stimulation of the superior colliculus and responsive to sensory inputs. Neurons activated by stimulation of the intermediate and deep collicular layers were distributed throughout the intralaminar thalamus. Approximately one half of them responded to tectal as well as sensory inputs. The majority were nociceptive or had a more complex response pattern including responses to auditory stimulation. A small population of low threshold units had contralateral orofacial receptive fields and responded to light taps; these units were preferentially localized anteriorly in the central lateral and paracentral nuclei. Neurons responsive to tectal and sensory stimulation were randomly intermingled with other neurons which had no detectable sensory input. The results indicate that ascending projection neurons of the intermediate and deep layers of the superior colliculus provide an input to functionally diverse subpopulations of intralaminar thalamic neurons. In view of its projections to motor cortex and basal ganglia, the intralaminar thalamus appears directly implicated in basal ganglia and superior colliculus related mechanisms of attention, arousal and postural orienting.

A PHA‐L analysis of ascending projections of the dorsal raphe nucleus in the rat

Ascending projections from the dorsal raphe nucleus (DR) were examined in the rat by using the anterograde anatomical tracer, Phaseolus vulgaris leucoagglutinin (PHA-L). The majority of labeled fibers from the DR ascended through the forebrain within the medial forebrain bundle. DR fibers were found to terminate heavily in several subcortical as well as cortical sites. The following subcortical nuclei receive dense projections from the DR: ventral regions of the midbrain central gray including the 'supraoculomotor central gray' region, the ventral tegmental area, the substantia nigra-pars compacta, midline and intralaminar nuclei of the thalamus including the posterior paraventricular, the parafascicular, reuniens, rhomboid, intermediodorsal/mediodorsal, and central medial thalamic nuclei, the central, lateral and basolateral nuclei of the amygdala, posteromedial regions of the striatum, the bed nucleus of the stria terminalis, the lateral septal nucleus, the lateral preoptic area, the substantia innominata, the magnocellular preoptic nucleus, the endopiriform nucleus, and the ventral pallidum. The following subcortical nuclei receive moderately dense projections from the DR: the median raphe nucleus, the midbrain reticular formation, the cuneiform/pedunculopontine tegmental area, the retrorubral nucleus, the supramammillary nucleus, the lateral hypothalamus, the paracentral and central lateral intralaminar nuclei of the thalamus, the globus pallidus, the medial preoptic area, the vertical and horizontal limbs of the diagonal band nuclei, the claustrum, the nucleus accumbens, and the olfactory tubercle. The piriform, insular and frontal cortices receive dense projections from the DR; the occipital, entorhinal, perirhinal, frontal orbital, anterior cingulate, and infralimbic cortices, as well as the hippocampal formation, receive moderately dense projections from the DR. Some notable differences were observed in projections from the caudal DR and the rostral DR. For example, the hippocampal formation receives moderately dense projections from the caudal DR and essentially none from the rostral DR. On the other hand, virtually all neocortical regions receive significantly denser projections from the rostral than from the caudal DR. The present results demonstrate that dorsal raphe fibers project significantly throughout widespread regions of the midbrain and forebrain.

GABAergic interneurons and neuropil of the intralaminar thalamus: an immunohistochemical study in the rat and the cat, with notes in the monkey

Immunohistochemistry using antibodies to glutamic acid decarboxylase (GAD) was used to investigate the intralaminar nuclei of the thalamus in rat, cat and monkey. Antibodies to gamma aminobutyric acid (GABA) were also used in the cat. Intralaminar immunoreactive cell bodies were not detected in the rat, but were clearly present in cat and monkey. In the latter species, GABA- or GAD-immunopositive perikarya were distributed throughout the anterior intralaminar nuclei, whereas in the posterior intralaminar complex they prevailed in the lateral part of the centre median nucleus and around the fasciculus retroflexus. Measurements of the area of immunostained intralaminar cell bodies in cat and monkey indicated that they are represented by small neurons. Experiments in the cat, based on retrograde tracers injections involving large sectors of the frontal and parietal cortices and the head of the caudate nucleus, revealed that the GABA- or GAD-immunoreactive cells and the retrogradely labeled projection neurons represented two separate intralaminar cell populations, although the latter also included small cells. Considerable differences were observed in the immunoreactive GABAergic neuropil of the anterior and posterior intralaminar nuclei. Clusters of densely packed bouton-like immunoreactive elements were detected in the former structures in the rat, cat and monkey, and were especially evident in the central lateral nucleus; immunopositive varicose fibers and puncta were diffusely distributed in the posterior intralaminar structures. Taken together with data from the literature, the present findings indicate that in cat and monkey local circuit inhibitory cells regulate not only the activity of principal thalamic nuclei which project densely upon restricted cortical fields, but also of the intralaminar structures which are widely connected with the cerebral cortex and the striatum. Regional variations in the distribution of GABAergic fibers and terminals suggest major differences in the organization of inhibitory circuits and synaptic arrangements of the anterior and posterior intralaminar thalamus.

Electrophysiological study of spinothalamic inputs to ventrolateral and adjacent thalamic nuclei of the cat

1. Extracellular and intracellular methods were used to record from fibers and neurons in the ventral lateral (VL) and adjacent nuclei of the cat thalamus. The receptive fields of the recorded units were analyzed and the units tested for inputs from the medial lemniscus (ML) and spinothalamic tract (STT) by electrical stimulation of the dorsal columns (DC) and ventrolateral funiculus (VLF) at the C2-3 spinal level. 2. Thirty-eight STT fibers were isolated in the thalamus. Their conduction velocities ranged from 15 to 75 m/s (mode 36 m/s). Adequate stimuli were found for 23 of these fibers. Seventeen were low-threshold (LT), 3 were wide-dynamic-range (WDR), and 3 were high-threshold (HT) units. 3. Five STT fibers were intra-axonally injected. Three were sufficiently well filled for analysis of their terminal fields. An intermediate-velocity STT fiber (conduction velocity 38 m/s) had a 4.3-microns axon and a single large terminal field in the central lateral nucleus (CL). The other two STT fibers were smaller, with diameters of 2.5 and 2.3 microns, conduction velocities of 15 and 19 m/s, and terminal fields made up of a few small boutons at the borders of the ventral posterior lateral nucleus (VPL). 4. Of 319 neurons isolated, 14 out of 129 (10.8%) in VL, 14 out of 76 (18.4%) in the VPL or ventral posterior medial (VPM) nucleus, 27 out of 64 (42.2%) in the CL nucleus, and 5 out of 50 (10%) in the reticular nucleus (R) responded at latencies less than 50 ms to VLF stimuli. A train of three pulses was more effective in driving VLF-responding neurons in all these nuclei than a single pulse. VLF-responding cells were widely dispersed in VL, concentrated in a focus in CL, and distributed around the borders of VPL. Most of those in VL and a small number in CL could be antidromically activated by stimulation of motor cortex. 5. Latencies of presynaptic responses (STT fibers) to VLF stimulation were short and varied from 0.8 to 3.9 ms (mode 1.6 ms). Despite this, very few fast-responding neurons were found. These were six VPL neurons (2.5 to 4 ms), one VL neuron (3 ms), and four CL neurons (3-4 ms). The initial spike latencies of the majority of thalamic neurons responding to VLF stimulation appeared in two peaks, one between 6 and 8 ms and the other at 10-15 ms.(ABSTRACT TRUNCATED AT 400 WORDS)

The amygdala in down's syndrome and familial Alzheimer's disease: Four clinicopathological case report

Senile plaques and neurofibrillary tangles were quantified in 14 subnuclei of the amygdala in the brains of 3 patients with Down's syndrome (DS), aged 19, 56, and 64 years, and in 1 patient with familial Alzheimer's disease (AD), aged 54 years. The amygdala of the 19-year-old Down's case contained numerous senile plaques (SPs) but no neurofibrillary tangles (NFTs). The distribution of neuropathological change in the amygdala was similar among the Down's and the Alzheimer's cases. Medial and ventral regions contained more SPs and NFTs than did lateral regions, and the SPs in ventromedial subnuclei generally were the “mature” type with a prominent amyloid core. In general, the numbers of SPs and NFTs were parallel in a given subnucleus with the striking exceptions of the deep medial basal, deep cortical, and lateral central nuclei that contained far more SPs than NFTs, and the medial and lateral superficial cortical nuclei that contained numerous NFTs but few SPs. Several subnuclei strongly interconnected with hippocampus and entorhinal cortex were more heavily involved than subnuclei related to the nucleus basalis of Meynert. The patterns of SP and NFT deposition are consistent with amygdaloid abnormalities found by others in sporadic AD. These findings demonstrate the similarity in amygdaloid pathology among Down's syndrome, familial Alzheimer's disease, and sporadic AD. The presence of senile plaques in the amygdala of the 19-year-old patient with DS suggests that the amygdala is a focus of early pathological change in DS and possibly AD.

Trigeminal nociceptive neurons in the subnucleus reticularis ventralis. II. Ascending projection

Ascending projection of trigeminal nociceptive neurons in the subnucleus reticularis ventralis (SRV) was studied in urethane/chloralose-anesthetized cats. Thalamic neurons having complex trigeminal receptive fields similar to those of SRV neurons were found in the intralaminar nuclei, i.e. nuclei centralis lateralis, centralis medialis and parafascicularis. About half of the SRV units tested were antidromically activated by electrical stimulation of the nucleus centralis lateralis. It was also found that a significant proportion of SRV units was antidromically activated by electrical stimulation of the mesencephalic reticular formation (MRF), and retrogradely labelled neurons were found in the SRV after HRP injection into the MRF. These findings suggest that SRV neurons relay trigeminal nociceptive inputs directly or indirectly via the MRF to the intralaminar nuclei.

Primate supplementary eye field. II. Comparative aspects of connections with the thalamus, corpus striatum, and related forebrain nuclei

The supplementary eye field (SEF) was defined electrophysiologically in behaving monkeys to study its connections with the diencephalon and corpus striatum. The specificity of SEF pathways was determined with horseradish peroxidase (HRP) histochemistry to compare its connections with those of the arcuate frontal eye field (FEF), contiguous dorsocaudal area 6 (6DC), and primary motor cortex (M1, arm/hand region). Results indicate that patterns of SEF connectivity were similar to the FEF and markedly different from areas 6DC and M1. Primary reciprocal thalamic pathways of the SEF were with the magnocellular ventral anterior (VA) nucleus, medial parvicellular VA, medial area X, and paralaminar medialis dorsalis (multiformis and parvicellularis). FEF showed similar connections but its most robust pathway was with MD rather than VA. In contrast, area 6DC showed the most extensive reciprocal connections with lateral VApc and lateral area X with only sparse connections with paralaminar MD. Area 6DC also exhibited reciprocal connections with the ventral lateral (VL) complex and the ventral posterior lateral nucleus, pars oralis (VPLo). M1 showed dense bidirectional connections with VPLo, and to a lesser extent, with VL. M1 pathways with the medial dorsal nucleus were negligible. All areas exhibited connections with the paracentral and central lateral nuclei and only M1 lacked connections with the central superior lateral nucleus. SEF and FEF exhibited similar efferent projections to the caudate and putamen. In the caudate, terminal fields were restricted to a central longitudinal core while those from area 6DC were more widely distributed. Eye field efferents were restricted to the putamen's face region while 6DC projections were more exuberant. The arm/hand region of M1 projected to the arm/hand region of the putamen. Pathways are discussed with respect to their significance in oculomotor control.

Spontaneous neuronal hyperactivity in the medial and intralaminar thalamic nuclei of patients with deafferentation pain

Electrical activity was recorded from single cells in the thalamus of 10 patients with chronic pain associated with deafferentation. Under local anesthesia, these patients underwent either electrode implantation or thalamotomy for treatment of their pain. In eight of the 10 patients, single units were identified as discharging spontaneously in high-frequency, often rhythmic, bursts. The discharges were of two types: short bursts comprised of two to six spikes with a burst frequency of one to four per second; and long trains of 30 to 80 spikes of similar frequency. Reconstruction of electrode trajectories indicated that recordings were made from the region corresponding to the lateral aspect of the mediodorsal thalamic nucleus, the central lateral nucleus, a small part of the central median nucleus, and the parafascicular nucleus. In the eight patients in whom spontaneous neuronal burst activity was exhibited, it was impossible to study activity evoked by natural cutaneous stimulation due to the continuous spontaneous neuronal discharges. Both animal and human studies have suggested that pain related to deafferentation is accompanied by spontaneous hyperactivity in the dorsal horn of the spinal cord and in the ventral posterior thalamic nuclei. The authors present evidence of spontaneous neuronal hyperactivity in the intralaminar thalamic nuclei of patients with pain related to deafferentation. The findings suggest that spontaneous neuronal discharge in patients with pain related to deafferentation is more widespread in the central nervous system than has been previously appreciated. The results have important implications for the surgical treatment of chronic pain.

Hippocampal pathway to the amygdala and stress ulcer development

The objective of the first study was to localize the reported aggravation of stress ulcers found after large bilateral hippocampal lesions in rats. Lesions in the ventral hippocampus produced a similar increase in the severity of gastric erosions after cold-restraint, as was seen after large bilateral lesions. Dorsal hippocampal damage produced no differential effects. In the second experiment, high-frequency electrical stimulation of the ventral CA1 region of the hippocampus, a procedure known to induce long-term potentiation, increased the evoked potentials in the lateral central nucleus of the amygdala, and in adjacent parts of the lateral and basolateral nuclei. The increase in the efficacy of synaptic transmission in this pathway attenuated stress ulcer development. It was concluded that the ventral hippocampus is part of a coping system, and a strengthening of synaptic connections with the central amygdala increases the coping ability of rats under stress conditions.

Organization of the thalamostriatal projections in the rat, with special emphasis on the ventral striatum

The organization of the thalamic projections to the ventral striatum in the rat was studied by placing injections of the retrograde tracer cholera toxin subunit B in the ventral striatum and small deposits of the anterograde tracer Phaseolus vulgaris-leucoagglutinin (PHA-L) in individual midline and intralaminar thalamic nuclei. In order to provide a complete map of the midline and intralaminar thalamostriatal projections, PHA-L injections were also made in those parts of the intralaminar nuclei that project to the dorsal striatum. The relationship of thalamic afferent fibres with the compartmental organization of the ventral striatum was assessed by combining PHA-L tracing and enkephalin immunohistochemistry. The various midline and intralaminar thalamic nuclei project to longitudinally oriented striatal sectors. The paraventricular thalamic nucleus sends most of its fibres to medial parts of the nucleus accumbens and the olfactory tubercle, whereas smaller contingents of fibres terminate in the lateral part of the nucleus accumbens and the most ventral, medial, and caudal parts of the caudate-putamen complex. The projections of the parataenial nucleus are directed towards central and ventral parts of the nucleus accumbens and intermediate mediolateral parts of the olfactory tubercle. The intermediodorsal nucleus projects to lateral parts of the nucleus accumbens and the olfactory tubercle and to ventral parts of the caudate-putamen. The projection of the rhomboid nucleus is restricted to the rostrolateral extreme of the striatum. A diffuse projection to the ventral striatum arises from neurons ventral and caudal to the nucleus reuniens rather than from cells inside the nucleus. Fibres from the central medial nucleus terminate centrally and dorsolaterally in the rostral part of the nucleus accumbens and medially in the caudate-putamen. Successively more lateral positions in the caudate-putamen are occupied by fibres from the paracentral and central lateral nuclei, respectively. The lateral part of the parafascicular nucleus projects to the most lateral part of the caudate-putamen, whereas projections from the medial part of this nucleus terminate in the medial part of the caudate-putamen and in the dorsolateral part of the nucleus accumbens. Furthermore, a rostral to caudal gradient in a midline or intralaminar nucleus corresponds to a dorsal to ventral and rostral to caudal gradient in the striatum. In the ventral striatum, thalamic afferent fibres in the "shell" region of the nucleus accumbens avoid areas of high cell density and weak enkephalin immunoreactivity.(ABSTRACT TRUNCATED AT 400 WORDS)

Neuronal cell types in the thalamic intralaminar central lateral nucleus of the cat

The existence of Golgi type II neurons was verified in the anterior intralaminar central lateral (CL) nucleus of the cat thalamus, and its projection cell types were identified, by means of Golgi impregnation. CL principal neurons were found to display a large- or medium-sized cell body and a radiate dendritic pattern. Their primary dendrites were limited in number, and had a rather long course; they were poorly ramified. The axons of principal neurons were impregnated only occasionally and for a short distance. Projection neurons of the 'bushy' or tufted type, described in the main thalamic sensory nuclei, were not identified in the CL in the present study. Typical Golgi type II neurons were found throughout CL. They were mainly small-sized, and displayed a rich dendritic arborization characterized by dendritic appendages. The axons of Golgi type II neurons were seen to give rise to extensive local arborizations. The present findings indicate that in the cat CL, principal cells are mainly represented by radiate neurons. Typical local circuit neurons also are evident in CL, suggesting that the activity of anterior intralaminar structures is regulated by intrinsic mechanisms similar to those operating in the main thalamic relay nuclei.

Preoccipital cortex receives a differential input from the frontal eye field and projects to the pretectal olivary nucleus and other visuomotor-related structures in the rhesus monkey

The bidirectional axonal transport capabilities of the horseradish peroxidase (HRP) technique facilitated the study of the frontal-eye-field (FEF) input and pretectal output of two regions of extrastriate preoccipital cortex (POC). Following horseradish peroxidase (HRP) gel implants into the middle and dorsal POC in two rhesus monkeys, the middle POC implant demonstrated retrograde frontal cortical labeling largely restricted to the inferior frontal eye field (iFEF) and adjacent inferior prefrontal convexity, whereas the dorsal POC implant showed labeling in the caudal ventral bank of the superior ramus of the arcuate sulcus (sas) and middle-to-dorsal region of the rostral bank of the concavity of the arcuate sulcus (dorsal FEF). Prominent anterogradely labeled efferent preoccipital projections were observed to the ipsilateral pretectal olivary nucleus (PON) and to a lesser extent the anterior pretectal nucleus. Although the middle POC case had heavier projections to the lateral PON, the dorsal case projected more heavily to the medial PON. In addition, both implants demonstrated subcortical connections with the lateral and dorsal inferior pulvinar nuclei, central superior lateral thalamic intralaminar nucleus, caudate nucleus, and middle-to-ventral claustrum. However, while the middle POC implant had efferent projections to the superficial superior colliculus (SC), pregeniculate nucleus (PGN), lateral terminal accessory optic nucleus (LTN), and dorsolateral pontine nucleus (DLPN), resembling those previously reported for the middle temporal (MT) visual area (Maunsell & Van Essen, 1982; Ungerleider et al., 1984), the dorsal implant had projections to the lateral intermediate SC, zona incerta (ZI), PGN, a notably lesser projection to the LTN, and basilar pontine projections to the lateral and lateral dorsal pontine subnuclei (not including the extreme dorsolateral DLPN). These preliminary results suggest that the preoccipital cortex, which reportedly functions in pupillary constriction, accommodation, and convergence, entertains connections with the PON and other visuomotor-related structures, and thus could act as an intermediary in the pathway between the iFEF and PON, and provide a possible explanation for pupillary effects that occur with stimulation of the FEF (Jampel, 1960) and within the contex of other oculomotor activities. The findings shed light on certain differences in connections of middle vs. dorsal POC with visuomotor-related nuclei, and appear to suggest that the middle region, which receives input from the iFEF, has greater access to the optokinetic (OKN) system by virtue of its projection to the LTN, and to the smooth-pursuit system b

Somatosensory neurons projecting from the superior colliculus to the intralaminar thalamus in the rat

Neurons of the rat superior colliculus projecting to the intralaminar thalamus were tested for their responses to somatosensory stimulation. They were identified by antidromic stimulation of the parafascicular nucleus and central lateral nucleus. To establish the existence of descending as well as ascending axon collaterals antidromic stimulation was applied to the upper cervical spinal cord in some cases. Somatosensory receptive fields were delineated and their laminar location in the superior colliculus was noted. Units were distributed throughout the intermediate and deep tectal layers, none were located in the superficial layers. Units with somatosensory receptive fields could be classified as low threshold, high threshold, wide dynamic range or complex. The majority of the peripherally responsive units (52%) were low threshold somatosensory units with contralateral receptive fields. All units were distributed throughout the intermediate and deep layers. Their distribution reflected the typical somatotopic organization of the superior colliculus. These results indicate that the intralaminar thalamus receives some sensory information by way of the tectum. In turn, the basal ganglia may gain direct access to this information by way of the thalamoneostriatal projection.

Distribution of ‘non-specific’ cholinesterase histochemical staining in the dorsal thalamus: a comparative study in rodents

Histochemical studies in rat dorsal thalamus demonstrate that ‘non-specific’ cholinesterase (ChE) enzyme activity is characteristic of neurons of the anterior dorsal (AD) and reuniens (Re) nuclei and in a cell group fouund as part of the central lateral (CL) and lateral dorsal (LD) nuclei. Extra-somatal ChE staining also is seen in the anterior ventral (AV) nucleus. Parallel histochemical studies in other rodents reveal slight ChE activity in neurons of the mouse AD and LD, but not in other thalamic nuclei. The dorsal thalami of hamsters, gerbils and guinea pigs show no detectable cellular staining of ChE, although low levels of extra-somatal ChE appear in AV and the internal medullary lamina. These data indicate that ‘non-specific’ cholinesterase activity is not found commonly in neurons of the dorsal thalamus and prominent ChE staining may be unique to the laboratory rat.

Supplementary eye field as defined by intracortical microstimulation: Connections in macaques

In macaques, the frontal eye field and the recently defined supplementary eye field play a role in the production of eye movements. Whereas the structure and function of the frontal eye field are well understood, little is known about the supplementary eye field. The goal of this study was to determine the connections of the physiologically defined supplementary eye field. In each case, the location of the supplementary eye field was determined by using intracortical microstimulation, the borders were marked with small electrolytic lesions, and horseradish peroxidase conjugated to wheat germ agglutinin was injected into the supplementary eye field. After the tissue was incubated with tetramethyl benzidine, it was determined that in three cases the injection site was confined to the physiologically defined supplementary eye field. The present results indicate that the supplementary eye field is reciprocally connected with the claustrum, ventral anterior nucleus, including pars magnocellularis, nucleus X, posterior subdivision of the ventral lateral nucleus, multiform, parvocellular, magnocellular, and densocellular subdivisions of the medial dorsal nucleus, central lateral nucleus, parafascicular nucleus, and suprageniculate-limitans complex. The supplementary eye field projects to the putamen, caudate, reticular nucleus of the thalamus, central densocellular nucleus, zona incerta, subthalamic nucleus, rostral interstitial nucleus of the medial longitudinal fasciculus, parvocellular part of the red nucleus, intermediate and deep layers of the superior colliculus, central gray, cuneiform nucleus, mesencephalic reticular formation, pontine gray, nucleus reticularis tegmenti pontis, and nucleus reticularis pontis oralis. The supplementary eye field is reciprocally and bilaterally connected with periprincipal and inferior prefrontal cortex, with periarcuate cortex, including the frontal eye field, the frontal ventral region, and with postarcuate premotor cortex, and cortex surrounding the supplementary eye field, including the supplementary motor area. The supplementary eye field is also reciprocally connected ipsilaterally with cortex in and around the cingulate sulcus and the intraparietal sulcus, whereas cortex within the superior temporal sulcus projects to the supplementary eye field. The connections of the physiologically defined supplementary eye field are compared to previously demonstrated connections of the supplementary motor region and of the physiologically defined frontal eye field. Comparisons between the connections of the frontal and supplementary eye fields reveal that both regions are connected with structures related to visuomotor functions, but the frontal eye field has more extensive connections with vision-related structures, and the supplementary eye field has more extensive connections with structures related to prefrontal and skeletomotor functions. Such connectional differences suggest functional differences between these two sensorimotor regions of the frontal lobe.

Properties of single neurons in the cat midsuprasylvian gyrus

Responses of cells in the midsuprasylvian gyrus (MSSG) of cats were investigated following electrical stimulation of the central lateral nucleus (CL) of the thalamus and tooth pulp, low-threshold cutaneous or visual afferents. Electrical stimulation in CL induced excitation in many cells located in cortical areas 5 and 7. Cells in these areas also received input from somato-sensory and visual afferents. Cells in MSSG showed a wide convergence from tooth pulp, low-threshold cutaneous afferents and from the CL. The majority of wide convergent cells in area 5 were found in layers IV and V, while cells excited by CL and tooth pulp were found in layers II and III. Similarities were found between CL and tooth pulp evoked responses with regard to the excitation-inhibition pattern. The excitation evoked from CL and tooth pulp was less often followed by a hyperpolarizing potential compared to that seen after low-threshold lip, paw and visual afferent stimulation. Stimulation sites in the lateral parts of CL-evoked responses with the shortest latencies in area 5. In this part of the cortex, short latency synaptic potentials were found in cells in superficial layers. In the same area, synaptic potentials of short latency were also evoked by electrical stimulation of tooth pulp, lip and paw. Light-flash stimulation evoked responses with the shortest latencies in area 7. The results of this study demonstrate that putative nociceptive information reaches the parietal association cortex and that part of this input may be relayed via CL. We suggest that the excitatory influences of nociceptive and CL stimulation is related to behavioral arousal and attention mechanisms.

Mesencephalic projections to the thalamic centralis lateralis and medial prefrontal cortex: a WGA-HRP study

In order to provide anatomical information for a possible pathway involved in pain mechanisms, rats were injected with horseradish peroxidase wheat germ agglutinin (WGA-HRP) in the centralis lateralis nucleus of the thalamus (Cl) or in the medial prefrontal cortex (PFCx) from which originated retrogradely labelled cells in the dorsal raphe nucleus (DR), locus cerules (LC) and surrounding structures. The locations of the Cl and the PFCx injections were previously determined by the presence of evoked single neuronal responses to noxious stimulations. The present study gives evidence for ascending pathways which originated in DR and LC and project to the Cl and PFCx. LC and DR projections suggest a possible route to an ascending modulation pain system.

Field potential analysis of the cortical projection of the central lateral nucleus in the cat.

A previous field potential study has indicated a monosynaptic projection of fibres from the central lateral nucleus (CL) to the mid-suprasylvian gyrus (MSSG). The present study, which is based on an analysis of current source density (CSD), aims to investigate further the sites of major localized synaptic activities in different layers of the MSSG after electrical stimulation in the CL. An initial positive surface potential was evoked in the MSSG with a latency of 3-5 ms and followed by a large negative potential with a peak latency of 8-15 ms. The initial positivity was only found in the rostral part of the MSSG, which corresponds to area 5. The positivity reversed in deeper layers. The CSD analysis showed a sink at a depth from 650 to 1050 microns. A corresponding source was found more superficially at 400-600 microns. This indicates that CL fibres have an excitatory synaptic termination on the soma or proximal dendrites of neurons in layers III and IV. The surface negative potential reversed at the border between layers II and III, suggesting a superficial CL projection. The CSD analysis of potentials in superficial layers showed a sink appearing between the pial surface and a depth of 350 microns, and a source lying in layers below. This indicates a depolarization of apical dendrites of cells in layers II and III. The superficial sink appeared in a large part of the MSSG. Application of a solution of 0.5% gamma-aminobutyric acid (GABA) on the surface of the cortex blocked the superficial sink and source and revealed a prominent sink current in layers III and IV in agreement with a deep termination of CL fibres. Application of a solution of 25 mM DL-2-amino-5-phosphono-valeric acid (APV) abolished CL-evoked cortical responses indicating that N-methyl-D-aspartate (NMDA) receptors are involved in the cortical activation. The CSD analysis confirms that CL has a wide superficial projection to the MSSG. It also confirms a deeper monosynaptic projection from CL to area 5.