Topographical arrangement of thalamic neurons projecting to the orbital gyrus in the cat

Distribution of thalamic neurons projecting to the orbital gyrus in the cat was examined by the horseradish peroxidase (HRP) method. After injection of HRP within the cortex of the orbital gyrus, thalamic neurons labeled with HRP were observed ipsilaterally within the pars parvicellularis of the nucleus ventralis posteromedialis (VPMpc) and the caudoventral portions of the nucleus centralis medialis: HRP-labeled neurons in the VPMpc were numerous and those in the nucleus centralis medialis were moderate in number. A few HRP-labeled neurons were seen occasionally in the nucleus paracentralis, nucleus centralis lateralis, nucleus submedius, and nucleus medialis. The VPMpc neurons labeled with HRP injected into the rostral portions of the orbital gyrus were seen chiefly in the dorsal aspects of the VPMpc, whereas the VPMpc neurons labeled with the enzyme injected into the caudal portions of the orbital gyrus were distributed mainly in the ventral aspects of the VPMpc: the region of distribution of the VPMpc neurons projecting onto the rostral portions of the orbital gyrus extended somewhat more rostrally than that of the VPMpc neurons projecting onto the caudal portions of the orbital gyrus. Functional significance of the VPMpc neurons that send their axons to the orbital gyrus is discussed in terms of the relay neurons of the central gustatory pathways.     

Peripheral input pathways projecting to the motor cortex in the cat

The possibility that the motor cortex receives peripheral input directly from the thalamus was examined using the evoked potential method and the following results were obtained. Potentials in the motor cortex evoked by stimulation of superficial radial (SR) or group II deep radial (DR) nerve were neither abolished nor delayed by ablation of the sensory cortex. Potentials in the motor cortex evoked by stimulation of group II DR nerve were most severely reduced by interruption of the spinocervcial tract. Potentials evoked by stimulation of SR nerve were more severely reduced in the sensory cortex than in the motor cortex by section of the dorsal funiculus or cooling of the cuneate nucleus. The size of evoked potentials in the motor cortex increased rapidly when stimulus intensity to DR nerve exceeded the threshold to group II fibers. The results suggest that some inputs from the SR and group II DR nerves reach the motor cortex without a relay through the sensory cortex.     

Distribution and size of thalamic neurons projecting to layer I of the auditory cortical fields of the cat compared to those projecting to layer IV

The distribution of thalamocortical neurons projecting to layer I of the cat auditory cortical fields was examined by the horseradish peroxidase (HRP) method. After HRP injection into layer I of the primary auditory cortex (AI), HRP-labeled neuronal cell bodies were distributed mainly in the medial, dorsal, and ventrolateral divisions of the medial geniculate nucleus (MGN) and suprageniculate nucleus (Sg), and additionally in the lateral and medial divisions of the posterior group of the thalamus (Pol and Pom), lateroposterior thalamic nucleus (Lp), and nucleus of the brachium of the inferior colliculus (BIN). After HRP injection into layer I of the second auditory cortex (AII), labeled neurons were seen mainly in the medial, dorsal, and ventrolateral divisions of the MGN and Sg and additionally in the Pom, Lp, and BIN. After HRP injection into layer I of the anterior auditory field (AAF), labeled neurons were located mainly in the medial and dorsal divisions of the MGN, Sg, Pol, and BIN, and additionally in the ventrolateral divisions of the MGN, Pom, and Lp. After HRP injection into layer I of the dorsal part of the posterior ectosylvian gyrus (Epd), labeled neurons were observed chiefly in the medial and dorsal divisions of the MGN, Sg, and Lp and additionally in the ventrolateral division of the MGN, Pom, and BIN. After HRP injection into layer I of the ventral part of the posterior ectosylvian gyrus (Epv), labeled neurons were distributed chiefly in the medial and dorsal divisions of the MGN and Pol and additionally in the ventrolateral division of the MGN, Sg, and BIN. Thus no labeled neurons were found in the ventral division of the MGN after HRP injection into layer I of all auditory cortical fields examined in the present study. The average soma diameters of neurons that were labeled after HRP injection into layer I were statistically smaller than those of neurons that were labeled after HRP injection into layer IV.     

Topographical organization of the thalamic afferent connections to the motor cortex in the cat

The topographical distribution of the cortical afferent connections to the different subdivisions of the motor cortex (MC) was studied in adult cats. The retrograde axonal transport of horseradish peroxidase technique was used. Small single injections of the enzyme were made in the entire MC, including the hidden regions in the depth of the sulcus cruciatus. The areal location and density of the subsequent thalamic neuronal labeling were evaluated in each case. Comparison of the results obtained in the various cases shows that the following: (1) The ventral anterior-ventral lateral complex is the principal thalamic source of afferents to the MC. (2) The ventral medial, dorsal medial, the different components of the posterior thalamic group (lateral, medial, and ventral posteroinferior and suprageniculate nuclei), and the intralaminar, lateral anterior, lateral intermediate, lateral medial, and anteromedial thalamic nuclei are also thalamic sites in which neural projections to the MC arise. (3) The thalamocortical projections to the MC are sequentially organized. The connections arising from the lateral part of the thalamus end in the region of area 4 that is situated medially in the superior lip of the sulcus cruciatus and in the posterior sigmoid gyrus. The projections originating in the most medial thalamic regions terminate in that region of area 6a beta which is located in the medial part of the inferior lip of the cruciate sulcus, and in the anterior sigmoid gyrus. Moreover, the ventral thalamic areas send connections to the most anteriorly located zones of the MC, while the most dorsal thalamic ones project to the most posteriorly located parts of the MC. (4) This shift in the thalamocortical connections is not restrained by cytoarchitectonic boundaries, either in the thalamus or in the cortex. (5) The populations of thalamocortical cells which project to neighboring MC subdivisions exhibit consistent overlapping among themselves. (6) These findings suggest, moreover, that the basal ganglia and the cerebellar projections to the MC through the thalamus are arranged in a number of parallel pathways, which may occasionally overlap.     

Receptive fields of motor neurons underlying local tactile reflexes in the locust

The receptive fields of motor neurons to a hind leg were mapped by recording intracellularly from their cell bodies or from the muscle fibers they innervate while stimulating mechanoreceptors on the surface of that leg. Each motor neuron is affected by a specific array of receptors that make up its receptive field. Boundaries along the anteroposterior or dorsoventral axes of the leg divide the receptive fields into excitatory and inhibitory regions. Proximodistal boundaries may correspond to the articulations between parts of the leg. Motor neurons that innervate antagonistic muscles have complementary receptive fields, so that the region that is excitatory for one is inhibitory for the other. The receptive fields of the motor neurons overlap. Tactile stimulation therefore leads to a specific local reflex that involves the coordinated movement of the segments of a leg. Five local reflexes are described, each of which moves the leg away from the site of stimulation. Afferents from the external mechanoreceptors do not synapse directly on the motor neurons, but instead on spiking local interneurons, some of which then synapse directly on motor neurons. These local interneurons have smaller receptive fields delineated by the same boundaries, so that the receptive fields of the motor neurons can be constructed from appropriate combinations of them. It is suggested that receptive fields are organized as "functional maps" that are appropriate for particular behavioral responses rather than solely to preserve or refine spatial information.     

Influence of dopamine on ventrolateral thalamic inputs in cat motor cortex

Neuronal activity in several brain regions is modulated by dopaminergic inputs. When single neuronal activity/20 trials of single-pulse ventrolateral thalamic (VL) stimulation was extracellularly recorded in the in vivo, anesthetized cat motor cortex, iontophoretic application of dopamine (DA) elicited either suppression or, in a fewer instances, facilitation of evoked unitary responses. The predominant inhibition exerted by DA appeared to be consistent for successive trials, and a D(1), D(2), and D(1)/D(2) receptor antagonist restored the effect, thereby reflecting a possible coexistence of two DA receptors. By contrast, only a fewer neurons' response to DA displayed facilitation, which was not attenuated by DA antagonists. Moreover, subsequent trials with receptor agonist and antagonists induced inconsistent effects. Except for the jitters, single unit spikes showed invariant latency, which was constant during all recording parameters, and the mean latency remained unchanged. The modulatory effects mediated by DA did not reveal any substantial difference between short- and long-latency responses. Both pyramidal tract neurons and non-pyramidal tract neurons, determined on the basis of antidromic potentials from the pyramidal tract, responded to DA essentially in a similar manner. It appears that DA overall inhibits cat motor cortical neuronal activity in response to VL inputs. We propose that such DAergic inhibition of thalamocortical excitation in the motor cortex could be critical for ongoing sensorimotor transformation.     

Quantitative analyses of thalamic and cortical origins of neurons projecting to the rostral and caudal forelimb motor areas in the cerebral cortex of rats

On the basis of studies using intracortical microstimulation, the existence of rostrocaudally separate two forelimb representation areas has been inferred in the motor cortex of rats. They are termed caudal and rostral forelimb areas (CFA and RFA). In this study, it was confirmed first that RFA and CFA are located in cytoarchitectonically distinct areas (medial and lateral parts of agranular cortex (AGm and AGl), respectively). In the second part of this study, the distribution of thalamic and cortical neurons projecting to RFA and CFA was quantitatively compared by injections of small and relatively constant amounts of retrograde fluorescent dyes (diamidino yellow and fast blue) into these areas. It was observed that (1) main inputs to RFA originated from AGl, namely CFA (2) CFA received dominant inputs from AGm including RFA and caudally adjacent granular cortex and (3) wider cortical areas and larger number of neurons projected to CFA than to RFA. As for the thalamocortical projections, both RFA and CFA received inputs from various thalamic nuclei, such as VL, VM, Po, PC, PF, CL, but cells projecting to RFA and CFA were differentially located in each nucleus. It was found that labeled cell number and/or density in VM, PC, CL, CM and MD after RFA injections were significantly larger than those after CFA injections. On the other hand, labeled cell number and/or density in VPL and VL were significantly higher after CFA injections than after RFA injections. In comparison with findings in primates, the results suggest that the cortical motor areas of rats may be specialized for different aspects of motor control.     

Sensory input to neurons of the motor area of the biceps in cat cortex

Correlation between activity of cortical neurons in the biceps motor area and a conditioning movement of the contralateral forelimb was studied in chronic experiments of two male cats. In the group of neurons related to the movement sensory inputs were not found in 68.1% of neurons, in the group not related to the movement in 97.9% of neurons. 24.2% of neurons from the first group had cutaneous receptive fields on the dorsal surface of the distal part of the moving forelimb, in this group 5 neurons had inputs from the forelimb joints.     

Quantitative analyses of neurons projecting to primary motor cortex zones controlling limb movements in the rat

The objective was to determine if projections of single neurons to primary motor cortex preferentially terminate in several efferent zones that could form synergies for the execution of limb movements. Intracortical microstimulation was used to identify zones evoking hip flexion (HF), elbow flexion (EF), and both plantarflexion (PF) and dorsiflexion (DF) about the ankle. Histological examination showed that the zones from which some movements were evoked extended beyond the agranular cortex into granular cortex. Fluorogold, Fast blue, and propridium iodide or rhodamine-labeled dextran were injected into three of these four efferent zones in each rat. There was a virtual absence of multiple-labeled cells despite having an intermingling of different-colored cells of which 15% in frontal cortex were less than 1.2 mm away from a neighboring neuron that projected to a different efferent zone. This suggests that single neurons projecting to the motor cortex do not hard-wire specific synergies but rather project to single efferent zones in order to offer the greatest degree of freedom for the generation of movements. The distribution of ventral posterolateral and ventrolateral thalamic nucleus labeling depended on whether the injections were in granular or agranular cortex. Conversely, frontal cortex projections to motor efferent zones were made irrespective of their location in either granular or agranular cortex and thereby supporting their presumed role in the control of movements. Hindlimb motor cortex injections yielded retrograde labeling that extended into the more localised distribution of frontal cortex neurons retrogradely labeled from forelimb injections. This may allow hindlimb movements to be synchronized by forelimb movements during walking on challenging terrain.     

Sources of thalamic afferent neurons, projecting into the suprasylvian gyrus of the dolphin cerebral cortex

The sources of a thalamic input to different loci of the suprasylvian gyrus (SSG) of the porpoise (Phocaena phocaena) cortex were studied by means of the retrograde HRP and fluorescent tracing methods. After injections of HRP into the anterior part of the SSG most cells were labelled in the lateral part of the ventrobasal complex. Some cells were also labelled in the ventroposteroinferior nucleus, posterior nucleus and caudally in the ventral parvocellular medial geniculate (MG). After injections of bisbenzimide in the middle part of the SSG many labelled cells were found in the ventral parvocellular MG and in the inferior pulvinar. Less cells were labelled in the magnocellular MG, lateral pulvinar and posterior nucleus. After bisbenzimide injection into the posterior part of the SSG the similar distribution of labelled cells was found but a sheet of labelled cells was shifted more laterally.     

Distribution of norepinephrine-containing neurons projecting to the parietal associative cortex and spinal cord in the cat locus coeruleus

The distribution of neurons projecting to the parietal associative cortex and spinal cord in the locus coeruleus (LC) of the cat was examined by the retrograde transport of the horseradish peroxidase (HRP) and catecholamine histofluorescence technique. It was demonstrated that neurons projecting to the parietal cortex were localized predominantly in the dorsal LC and the most dense labeling was observed at the coronal section corresponding to frontal plane P-1. Neurons projecting to the spinal cord were concentrated in the ventral LC and the largest quantity of labeled neurons was found in the frontal plane P-3. Microinjections of HRP into the parietal cortex and spinal cord also caused intense retrograde labeling of neurons in the reticular formations of mesencephalon, pons varolii and medulla oblongata. Coeruleo-cortical and coeruleo-spinal projection neurons formed in the LC two partially overlapping cellular populations which could not be differentiated according to morphologic characteristics of their cells. It was concluded that the parietal associative cortex of cat as well as spinal cord had direct afferent inputs both from the LC and from the reticular formation.     

The inhibitory input from the substantia nigra to the mediodorsal nucleus neurons projecting to the prefrontal cortex in the cat

The effects of stimulation of the substantia nigra pars reticulata (SNr) on the neurons of the mediodorsal nucleus (MD) projecting to the prefrontal cortex (PF) were studied in cats. The MD neurons projecting to the ventral part of the PF tended to be located in the ventral part of the MD, while those projecting to the dorsal part of the PF in the dorsal part. Spontaneous discharges of 30/57 tested MD neurons were suppressed by SNr stimulation at a latency ranging from 2 to 15 ms. The latency of the suppression corresponded well to that of antidromic responses of SNr neurons elicited by MD stimulation (from 1.4 to 14.0 ms). Intracellular recordings in a few MD neurons showed IPSP by SNr stimulation. The SNr is considered to exert an inhibitory effect on the MD neurons projecting to the PF.     

Morphological and functional properties of trigeminal nucleus oralis neurons projecting to the trigeminal motor nucleus of the cat

Horseradish peroxidase (HRP) was injected into the somata located in the rostrodorsomedial part (Vo.r) of the trigeminal nucleus oralis; an axonal projection to the trigeminal motor nucleus (Vmo) was demonstrated in two Vo.r neurons. The two neurons differed in their morphological and functional properties. The first Vo.r neuron responded to stimulation of low-threshold mechanoreceptors and its stem axon gave off massive axon collaterals that issued terminal branches to the dorsolateral subdivision of Vmo, Vo.r, and the medial and lateral parts of the lower brainstem reticular formation. The second Vo.r neuron was activated by stimulation of the tooth pulp or lingual nerve at twice longer latency than that of the first neuron. This stem axon was divided into two main ascending and one descending branches, and one of the main ascending branches was further bifurcated into two branches. The main non-bifurcated ascending branch gave off 4 collaterals, two of which sent terminal branches into the dorsolateral subdivision of Vmo and others into the Vo.r and juxta-trigeminal regions. The somato-dendroarchitectonic differences were also described in the two Vo.r neurons stained.     

Response properties and morphological identification of neurons in the cat motor cortex

Intracellular recordings and morphological identification of neurons using intracellular HRP staining were performed in the cat motor cortex. By thalamic ventrolateral (VL) or cerebellar nucleus stimulation, pyramidal cells in layer III, fast pyramidal tract neurons (PTNs) and stellate cells in layers II and III were activated with short latency and fast rising EPSPs, while pyramidal cells in layer II and slow PTNs showed longer latency and slow rising EPSPs. This difference may be related to activation through the deep and superficial thalamocortical projections. Although pyramidal cells in layer VI did not respond orthodromically to VL or cerebellar stimulation, some of them proved to receive the recurrent action of PTNs because of the response to stimulation of the cerebral peduncle (CP). One aspinous stellate cell in layer III was activated by CP as well as VL stimulation. This cell was supposed to be an inhibitory interneuron responsible for both recurrent and VL-evoked inhibition.     

Receptive fields of neurons in areas 3b and 1 of somatosensory cortex in monkeys

Receptive fields of neurons within the separate representations of the glabrous hand in areas 3b and 1 of somatosensory cortex were studied in cynomolgus monkeys. Many neurons in area 1 have center-surround receptive fields with separate 'on' and 'off' zones, while neurons in area 3b exhibit largely uniform or homogeneous receptive fields.     

Neuronal populations in the posterior group of thalamic nuclei projecting into the amygdaloid complex and auditory cortex in the cat

Location of neurons in posterior thalamic nuclei and neighbouring structures of the midbrain regions projecting to the amygdaloid complex and auditory cortex of cat was studied by the method of horseradish peroxidase. The main sources of these brain region projections to amygdaloid complex are peripeduncular , subparafascicular and suprageniculate nuclei and caudal division of the medial geniculate body. The cells of origin of projections to the auditory cortex are located in all medial geniculate nuclei and wide regions of the posterior thalamic group. Neuron pools projecting to the auditory cortex and amygdala exist in medial parts of the posterior thalamic nuclei. The role of posterior thalamic nuclei in transmission of auditory signals to amygdala is discussed.