The callosal connections of the primary somatosensory cortex and the neural bases of midline fusion.
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Action Potentials↗
Biomedical subjects
T Manzoni
Publications and source records attributed to T Manzoni.
Immunocytochemical evidence for glutamatergic cortico-cortical connections in monkeys.
Retrograde transport of horseradish peroxidase and immunocytochemical visualization of glutamate (Glu) were combined to investigate the neurotransmitter used by cortico-cortical neurons in the first (SI) and second (SII) somatic sensory areas of macaque monkeys. The majority of association and callosal neurons in SI and SII were immunoreactive for an antiGlu serum: evidence was therefore obtained in support of a role for Glu, or a closely related compound, as the synaptic transmitter used by cortico-cortical fibers.
Animals↗
Demonstration of glutamate-positive axon terminals forming asymmetric synapses in cat neocortex.
Electron microscopic examination of sections immunocytochemically processed with an anti-glutamate serum reveals that many asymmetric synapses in the cat neocortex contain elevated levels of immunodetectable glutamate. These labelled axon terminals are likely to use glutamate as neurotransmitter. Axon terminals forming symmetric contacts were never labelled. Since glutamate is known to exert potent excitatory effects on neocortical neurons, the present finding gives immunocytochemical evidence that asymmetric synapses are excitatory.
Animals↗
Glutamate-positive corticocortical neurons in the somatic sensory areas I and II of cats.
Combined retrograde transport-immunocytochemical experiments were carried out on cats to study the morphology, laminar distribution, and percentages of corticocortical projecting neurons of somatosensory area I (SI) and II (SII) showing immunoreactivity to an antiserum raised against the amino acid glutamate (Glu). A previously characterized anti-Glu serum (Conti et al., 1987a, b; Hepler et al., 1987) was used in conjunction with HRP. This tracer was injected either in SI to label retrogradely neurons in ipsilateral SII (SII-SI association neurons) and contralateral SI (SI-SI callosal neurons) or in SII to label retrogradely neurons in ipsilateral SI (SI-SII association neurons) and contralateral SII (SII-SII callosal neurons). In sections from SI and SII processed for simultaneous visualization of Glu and HRP (Bowker et al., 1982), and containing the cells from which every one of the 4 corticocortical projections arise, 3 types of labeled neurons were observed: (1) single-labeled neurons showing the homogeneous brown immunoreaction product of Glu (Glu-positive neurons); (2) single-labeled neurons containing the granular black reaction product of retrogradely transported HRP (Glu-negative, association or callosal neurons); and (3) double-labeled neurons in which both the black HRP granules and the brown immunostaining were present (Glu-positive, association or callosal neurons). Double-labeled neurons were all pyramidal in shape and were distributed intermingled with Glu-negative corticocortical neurons in all layers of SI and SII known to give rise to association and callosal projections. Counts from 25-micron-thick sections showed that of 432 association and callosal neurons sampled from SI and SII, 214 (49.5%) were Glu-negative and 218 (50.5%) Glu-positive. In counts carried out on 5-micron-thick sections, the percentage of Glu-positive corticocortical neurons raised to about 70%. The 2 populations of single- and double-labeled corticocortical neurons showed no difference in their perikaryal cross-sectional areas. The present results show that a large fraction of association and callosal neurons of SI and SII are immunoreactive for Glu, and, therefore, these neurons probably use this excitatory amino acid, or a closely related compound, as neurotransmitter.
Animals↗
D-[3H]aspartate retrograde labelling of callosal and association neurones of somatosensory areas I and II of cats.
Experiments were carried out on cats to ascertain whether corticocortical neurones of somatosensory areas I (SI) and II (SII) could be labelled by retrograde axonal transport of D-[3H]aspartate (D-[3H]Asp). This tritiated enantiomer of the amino acid aspartate is (1) taken up selectively by axon terminals of neurones releasing aspartate and/or glutamate as excitatory neurotransmitter, (2) retrogradely transported and accumulated in perikarya, (3) not metabolized, and (4) visualized by autoradiography. A solution of D-[3H]Asp was injected in eight cats in the trunk and forelimb zones of SI (two cats) or in the forelimb zone of SII (six cats). In order to compare the labelling patterns obtained with D-[3H]Asp with those resulting after injection of a nonselective neuronal tracer, horseradish peroxidase (HRP) was delivered mixed with the radioactive tracer in seven of the eight cats. Furthermore, six additional animals received HRP injections in SI (three cats; trunk and forelimb zones) or SII (three cats; forelimb zone). D-[3H]Asp retrograde labelling of perikarya was absent from the ipsilateral thalamus of all cats injected with the radioactive tracer but a dense terminal plexus of anterogradely labelled corticothalamic fibres from SI and SII was observed, overlapping the distribution area of thalamocortical neurones retrogradely labelled with HRP from the same areas. D-[3H]Asp-labelled neurones were present in ipsilateral SII (SII-SI association neurones) in cats injected in SI. In these animals a bundle of radioactive fibres was observed in the rostral portion of the corpus callosum entering the contralateral hemisphere. There, neurones retrogradely labelled with silver grains were present in SI (SI-SI callosal neurones). Association and callosal neurones labelled from SI showed a topographical distribution similar to that of neurones retrogradely labelled with HRP. The laminar patterns of corticocortical neurones labelled with D-[3H]Asp or with HRP were also similar, with one exception. In the inner half of layer II, SII-SI association neurones and SI-SI callosal neurones labelled with the radioactive marker were much less numerous than those labelled with HRP. In cats injected in SII, D-[3H]Asp retrogradely labelled cells were present in ipsilateral SI (SI-SII association neurones). Their topographical and laminar distribution overlapped that of neurones labelled with HRP but, as in cats injected in SI, association neurones labelled with silver grains were unusually rare in the inner layer III.(ABSTRACT TRUNCATED AT 400 WORDS)
Animals↗
Callosal projections from area SII to SI in monkeys: anatomical organization and comparison with association projections.
The present research was aimed at ascertaining in the macaque monkey the reciprocity of the heterotopical callosal connections between SI and SII, with particular regard to the connectivity of the hand representation, and at comparing the topographical and laminar pattern of these callosal connections with those of association connections entertained by these areas. Horseradish peroxidase (HRP) was unilaterally injected into area SI in five monkeys. The sites of HRP delivery included the trunk and the hand zones preliminarily identified by recording multi-unit responses to peripheral stimulation by means of microelectrodes. Anterograde and retrograde labelling was studied in SII of both sides. The results showed the complete reciprocity of the heterotopical callosal connections between SI and SII. In the latter area both callosal axon terminals and neurones were found, which were labelled from either the trunk or the hand zone of contralateral SI. Labelling of callosal axon terminals occurred mainly in layer IV and in the lowermost part of layer III. Labelled callosal neurones were mainly in the lower half of layer III, whereas few occurred in infragranular layers. Topographically, the distribution of callosal terminals and cell bodies duplicated the distribution of association terminals and cell bodies labelled in SII on the side ipsilateral to HRP injection. The laminar pattern of termination of association fibres from SI was similar to that of callosal fibres. However, the distribution of association-projecting neurones in SII showed a striking difference from that of callosal-projecting neurones. Unlike the latter neurones, which were mainly located in supragranular layers, association cell bodies overwhelmingly dwelt in layers V and VI and were less numerous in layers II and III. This laminar pattern of association SII-SI cells corresponds to the "feed-backward" model and fits the laminar pattern of their axon terminations (Friedman: Brain Res. 273: 147-151, '83). The association and callosal inputs and outputs of area SII are discussed in relation to the function of the forward and backward type of reciprocal connections entertained with SI in the ipsilateral hemisphere and to the function of SII in the interhemispheric exchange of somatosensory information.
Animals↗
D-[3H]aspartate retrograde labelling of association neurones in area SI of the cat.
Injection of D-[3H]aspartate into area SII of cats retrogradely labelled association cells in area SI. Numerous intensely labelled cells were found in layer II and in the upper layer III but labelling was scanty in other layers. In contrast, association neurones labelled by horseradish peroxidase injected in area SII mixed with the radioactive marker were also numerous in the other sublaminae of layer III and in infragranular layers of SI. Association neurones of the outer laminae of this area are likely to use aspartate and/or glutamate as neurotransmitter(s).
Animals↗
Bilateral receptive fields and callosal connectivity of the body midline representation in the first somatosensory area of primates.
Experiments were performed in order to study the receptive field (RF) organization and the callosal connectivity of the trunk representation zone in areas 3b and 1 of the postcentral cortex of macaque monkeys. Multiunit microelectrode recordings showed that neurons responding to tactile stimulation of bilateral RFs across the midline of the body were contained in three topographically distinct zones of the trunk map. In one zone, at the junction between cytoarchitectonic areas 3b and 1, RFs straddled the dorsal midline of the trunk. In the other two zones, one located caudally in area 1 in front of the postcentral dimple, and the other rostrally in area 3b in the depth of the posterior bank of the central sulcus, RFs straddled the ventral midline of the trunk. The first one and the other two zones are referred to here as the dorsal and the ventral midline representation zones, respectively. Elsewhere in the trunk map, neurons responded only to stimulation of contralateral RFs. The callosal connectivity of the trunk map was studied by means of the transport of horseradish peroxidase (HRP). Multiple injections of HRP in electrophysiologically identified sites of the trunk representation in one hemisphere labeled both callosal fiber terminals and callosally projecting neurons in the contralateral homotopic cortex. Dense patches of callosal neurons intensely labeled with HRP were present in the cortical regions representing the body midlines and were distributed for the most part in layer III. Some neurons lightly labeled with HRP were scattered in other zones of the trunk map. Callosal terminations were densest within the midline zones and very sparse or absent in the lateral trunk zones. Correlation of physiological and anatomical data obtained either separately or from the same animal demonstrated that cortical regions containing bilateral-field neurons also contained the highest density of labeled callosal terminations and neurons. This correlation suggests a role for the corpus callosum in the perception of the body midline, either by generating the bilateral RFs of these neurons or by coordinating the activity of the regions containing neurons with thalamically generated bilateral RFs.
Abdomen↗
[Topography of the thalamo-cortical projections on trunk representation demonstrated by fluorescent neuro-tracers].
With the aim to study the detailed topography of the thalamo-cortical neurones projecting to the trunk representation zone of the first somatosensory area (SI), punctate injections of three different fluorescent tracers (Evans Blue, Nuclear Yellow and Fast Blue) were performed in the three physiologically defined subareas forming the trunk region of SI. These injections resulted in the labelling of three different cell aggregates, narrow in dorsoventral and mediolateral extent but elongated rostrocaudally, located in topographically distinct regions of the nucleus ventralis posterio-lateralis. The results suggest that the highly organized topography of the trunk representation of area SI is imposed by the thalamo-cortical input from VPL.
Amidines↗
[Callosal connections of the somatosensory area in the primate: anatomical and electrophysiological studies].
In order to study the callosal connections of the hand sensory field of the second somatosensory area of the monkey, experiments were carried out by combining the method of retrograde neuronal tracing with microelectrode recording. In six monkeys, Macaca Irus, single or multiple (5-8) injections of horseradish peroxidase (HRP) were performed into the cortex of the parietal operculum of one side. Neurones retrogradely labelled with HRP (callosal neurones) were found in the post-central gyrus and in the parietal operculum of the contralateral hemisphere. Microelectrode recording from this hemisphere showed that the cortical zones of both the first and the second somatosensory area containing neurones excited by sensory stimulation of the contralateral hand also contained HRP-positive neurones.
Animals↗
Topography and receptive field organization of the body midline representation in the ventrobasal complex of the cat.
The topography and receptive field (RF) organization of neurones in the trunk zone of the thalamic ventrobasal complex (VB) projecting to the homologous zone of the ipsilateral first somatosensory area (SI) were studied in the cat by performing experiments of retrograde neuronal tracing and microelectrode recording. Punctate cortical injections of small amounts of either horseradish peroxidase or fluorescent tracers (Evans Blue, Nuclear Yellow and Fast Blue) retrogradely labelled cell aggregates lying in the dorsal half of a VB region interposed between subnucleus VPL1 and VPLm. Aggregates of labelled cells were narrow in dorsoventral and mediolateral extent and elongated rostrocaudally. The distribution of VB cells projecting to the cortical subareas representing the dorsal midline, lateral trunk and ventral midline of the body in area SI, was established by injecting a different fluorescent marker into a physiologically defined site in each subarea. These injections resulted in labelling of three different cell aggregates located in topographically distinct regions of the VB trunk zone. Each aggregate of labelled cells only projected to one cortical subarea. Microelectrode analysis of cell populations of the VB trunk zone showed that neurones lying in regions projecting to dorsal and ventral midline zones of area SI had bilateral RFs, straddling the dorsal and the ventral midline of the body respectively. Neurones lying in the region projecting to the lateral trunk representation of area SI had contralateral RFs located on the lateral surface of the trunk. The results suggest that the detailed topography of the trunk map in the area SI and the bilaterality of the cortical representation of the body midlines, described in previous experiments, is imposed by the thalamocortical input from the VB.
Animals↗
Callosal mechanism for the interhemispheric transfer of hand somatosensory information in the monkey.
The retrograde transport of horseradish peroxidase (HRP) was combined with extracellular microelectrode recording from single and multiple-neurones to study the anatomical and functional organization of the callosal connections of the hand sensory projection field in the parietal operculum of monkeys (Macaca Irus). In 3 animals anaesthetized with ketamine, a single injection of HRP (0.5 microliter) was delivered into the cortex forming the upper bank of the sylvian sulcus at a site where neuronal responses to somatic sensory stimulation of the hand were recorded. In the ipsilateral hemisphere, retrogradely HRP-labelled cells were found in the cortex of the post-central gyrus and in the thalamic nuclei ventralis posteroinferior and pulvinar oralis. In the contralateral hemisphere HRP-labelled neurones were present in the opercular cortex lying dorsal, and slightly caudal, to the posterior pole of the insula. Few scattered callosal neurones were also found in the post-central gyrus. In 3 other animals, multiple injections (5-8; 0.5 microliter each) of HRP were performed in the parietal operculum. In the ipsilateral hemisphere, retrogradely labelled cells were present in the post-central gyrus and in the following thalamic nuclei: ventralis posteroinferior, pulvinar oralis and medialis, ventralis posteromedialis and posterior complex. Few labelled cells were also present in the ventral part of the nucleus ventralis posterolateralis. In the contralateral hemisphere, numerous callosal cells were labelled with HRP. These cells were found, with regional variations in density, in wide regions of the buried and exposed cortex of the parietal operculum and in the post-central gyrus. These 3 monkeys were subjected to microelectrode mapping experiments (N2O and halothane anaesthesia) to explore the peripheral receptive fields of neurones in the parietal operculum and post-central gyrus contralateral to the injected side. HRP labelled callosal neurones were found in regions of the second and first somatosensory cortical areas which also contained units driven from the contralateral hand.
Animals↗
Axonal branching in the periaqueductal gray projections to the thalamus: a fluorescent retrograde double-labeling study in the cat.
The double-labeling technique based on the retrograde axonal transport of fluorescent tracers (Evans blue, EB; Fast blue, FB; Nuclear yellow, NY) was used in the cat in order to investigate the occurrence of axonal branching in the periaqueductal gray (PAG) projections to some thalamic nuclei (n. ventralis postero-lateralis, VPL; n. ventralis postero-medialis, VPM; n. parafascicularis, Pf). In a first group of cats, FB and EB were injected, respectively, within the right and left VPM. In another two groups of cats, FB injections into Pf were combined with either EB or NY injections within VPL or VPM. Double-labeled neurons were found within the PAG only in the animals of the first group. The present results show that some PAG neurons project bilaterally to VPM by means of axons collaterals.
Animals↗
Cortico-cortical neurones of somesthetic area SI as studied in the cat with fluorescent retrograde double-labelling.
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Animals↗
Periaqueductal grey projection to the ventrobasal complex in the cat: an horseradish peroxidase study.
Horseradish peroxidase (HRP) was injected within the thalamic ventrobasal complex of 14 cats. The aim was to ascertain whether the periaqueductal grey matter (PAG) sends fibres to this complex. Retrogradely labelled cells were found within the PAG following HRP delivery either in the nucleus ventralis posterolateralis (VPL) or ventralis posteromedialis (VPM). PAG-VPL projection is only ipsilateral and arises mainly from lateral PAG, PAG-VPM projection is bilateral and originates from latero-ventral regions of the central grey. The hypothesis that PAG might control the activity of ventrobasal nociceptive neurones is proposed.
Afferent Pathways↗
Callosal projections from the two body midlines.
1. Horseradish peroxidase (HRP) was injected within the proximal limb and trunk representation zones of the first somatosensory area (SI) of 16 cats. The tangential and laminar distributions of retrogradely labelled neurones (callosal neurones) of the contralateral homotopic cortex were studied. This cortex was explored with microelectrodes on the day after HRP delivery to relate the distribution of callosal neurones to the electrophysiological map of the trunk. 2. Callosal neurones were found in the contralateral SI area mainly in layer III, but also many in layer VI, especially following large HRP injections, and very few in the other layers. Callosal neurones of layer III were mostly pyramidal, those of layer VI pyramidal and non-pyramidal. Many neurones were intensely stained by HRP, and cell details, such as fine dendritic branchings, spines, and axon collaterals, could be seen. 3. Callosal neurones are grouped within two regions located, respectively, in the rostral and caudal parts of the exteroceptive trunk map. The rostral region overlaps the representation of the dorsal midline (cytoarchitectonic field 3b) and the second one that of the ventral body midline (cytoarchitectonic field 2). The cortex intermediate between these two fields contains rare callosal cells and receives afferences from the lateral trunk surface. Few or no callosal cells were found within the proximal limb zones. Neurones recorded from the two midline zones have bilateral receptive fields straddling either the back or the ventral surface of the trunk. 4. It is concluded that the interhemispheric fusion between the two hemibody representations in areas SI is brought about by the mutual callosal links which the two midline zones entertain with their contralateral homologues.
Animals↗
The anatomical substrate of callosal messages from SI and SII in the cat.
Horseradish peroxidase (HRP) was injected into the first (SI) or second (SII) somatosensory areas of 21 adult cats. The radial and tangential (normal and parallel to the pial surface, respectively) distribution and morphology of the callosal neurons were studied. HRP injections were combined with single unit recording in the contralateral cortex in order to determine which part of the somatosensory periphery is represented within the regions containing callosal neurons, the callosal (efferent) zones, in SI and SII. The callosal zone of SI extends over the trunk and part of the forepaw representation. In the forepaw and hindlimb representations callosal neurons projecting only to the contralateral SII are found, while in the trunk representation callosal neurons projecting to contralateral SI or SII are found. The callosal zone in SII extends widely throughout the forepaw representation in this area and projects to the contralateral SII but not to SI. In both SI and SII the callosal neurons are mainly located in layer III. A few of them are also found in layer VI. They are very rare in other layers. Callosal neurons in layer III are mostly pyramidal but exceptionally stellate; in layer VI they are pyramidal, triangular, and occasionally stellate. These data indicate that transformations of the cortical somatosensory maps are achieved in the message sent through the corpus callosum. These transformations are i) determined by the extent and location of the callosal zones and perhaps by the distribution of callosal neurons within them, ii) different in different areas, iii) different in a same area, according to the cortical targets to which they are conveyed. The existence of callosal connections originated from areas of distal forepaw representation supplies a possible anatomical substrate for those types of intermanual transfer of tactile learning which depend upon the integrity of the corpus callosum.
Animals↗
Anatomical and functional aspects of the associative projections from somatic area SI to SII.
1. Electrophysiological and morphological (retrograde axonal transport of horseradish peroxidase, HRP) experiments have been carried out in the cat in order to study the associative projections from area SI to ipsilateral SII. 2. Microelectrode recordings were performed in the forepaw focus of SII both in normal (64 units) and in SI-undercut (51 units) cats. 29.6% of the neurons recorded in the unoperated and 29.4% of those collected in the operated cats were excited by electric stimulation of the ipsilateral SI (forepaw focus). In both preparations almost all such units were endowed with large (either contra- or bilateral) receptive fields (RF). Cell population recorded in the SI-undercut cats showed no significant impairment to peripheral stimuli and/or changes in the size of the RFs. 3. From the forepaw focus of SI, 150 units have been recorded and tested by stimulation of the homologous focus of the ipsilateral SII. Eight of them were fired antidromically and thus identified as association cells. Their RFs were very small and located only in the digits of the contralateral forepaw. 4. Both single or multiple HRP injections were performed in SII. Retrogradely labelled cells were found in the ipsilateral SI. The great majority of association cells are pyramids and dwell mainly in layer III. In spite of the large diffusion of the exogenous reaction product in the injected SII and of the presence of retrogradely labelled cells anywhere in the ipsilateral thalamic VB complex, the distribution of association cells is unequal throughout SI since they strongly predominate in the digit zone of the forepaw representation.
Animals↗