Emergence of callosally projecting neurons with stellate morphology in the visual cortex of the kitten

Summary Callosally projecting neurons in areas 17 and 18 of the adult cat can be classified into two types on the basis of their dendritic morphology: pyramidal and stellate cells. The latter are nearly exclusively of the spinous type and are predominantly located in upper layer IV. Retrograde transport of the carbocyanine dye DiI, applied to the corpus callosum, showed that, up to P6, all callosally projecting neurons resemble pyramids in the possession of an apical dendrite reaching layer I. At P10, however, callosally projecting neurons with stellate morphology were found. A study was designed to distinguish whether these neurons are late in extending their axons to the corpus callosum or, alternatively, have transient apical dendrites. To this end, callosally projecting neurons were retrogradely labeled by fluorescent beads injected in areas 17 and 18 at P1–P3 and then either relabeled with DiI applied to the corpus callosum at P10 or intracellularly injected with Lucifer Yellow at P57. Double-labeled stellate and pyramidal cells were found in similar proportions to those found for the total, single-labeled population of callosally projecting neurons. It is therefore concluded that callosally projecting spiny stellate cells initially possess an apical dendrite and a pyramidal morphology. At P6, i.e. close to the time when stellate cells appear, layer IV neurons with an atrophic apical dendrite were found, suggestive of an apical dendrite in the process of being eliminated.     

A direct pathway from thalamus to visual callosal neurons in cat

Summary Horseradish peroxidase was injected in the right visual cortex and a large electrolytic lesion made in the left lateral geniculate nucleus of an adult cat. Neurons of origin of the callosal projection to the injected cortex were identified by retrograde labelling and selected for electron microscopic study. Degenerating thalamo-cortical axon terminals were found to contact a labelled stellate cell in layer IV and a labelled pyramidal cell in layer III at the border region of areas 17 and 18. We conclude that there is a monosynaptic pathway from lateral geniculate nucleus to the cells of origin of callosal axons to the contralateral visual cortex.Supported by the Swiss National Science Foundation (3.0950.77)     

The study of Golgi stained cells and of experimental degeneration under the electron microscope: a direct method for the identification in the visual cortex of three successive links in a neuron chain.

A direct method is presented which makes it possible to identify, from synapse to synapse, three successive links of a neuron chain. The potentialities of the method are shown by examples of the termination of specific afferents in the visual cortex. Following unilateral lesion of the lateral geniculate nucleus in the rat, the distribution of degenerating geniculocortical boutons was studied on two Golgi-stained cells in layer IV of the primary visual cortex. One of the cells was definitely a small pyramidal cell; the other was identified as a spiny stellate (although the possibility that it too was a small pyramidal cell was not rigorously excluded). Both cells received monosynaptic input from the specific afferents as proved by the existence of degenerating boutons synapsing on their dendritic spines. The axonal arborizations of both Golgi-stained cells were traced at the electron microscopic level in thin section series in order to identify the postsynaptic structures contacted by their boutons. All boutons studied established asymmetrical contacts and about 50% of the synapses given by the impregnated boutons were found on smooth dendritic shafts of stellate cells, the rest on spines. The results obtained suggest a neuron circuit involving, successively, the visual afferents, spiny interneurons or monosynaptic visual target pyramidal cells and nonspiny stellate cells.It is suggested that a similar approach might provide direct information about the connectivity in neuron networks in many other parts of the central nervous system hitherto defying elucidation with conventional methods.     

Morphology of identified entorhinal neurons projecting to the hippocampus. A light microscopical study combining retrograde tracing and intracellular injection

Cells of origin of the entorhinohippocampal pathway were retrogradely labeled by injection of Fast Blue into the ipsilateral hippocampus. The cells, which were located in layers I, II and III of the lateral entorhinal cortex, were then intracellularly injected with Lucifer Yellow to reveal their complete morphology. We could thus establish that among the hippocampally projecting entorhinal cells there are pyramidal and pyramid-like cells, spiny stellate cells of various shapes, sparsely spinous horizontal and multipolar cells. The involvement of horizontal and multipolar neurons in projections has not previously been recognized although all of these cell types have already been described in Golgi studies.

The results indicate that the organization of the perforant path is more complex than has been assumed. Finally, they are at variance with the classical concept which subdivides cortical neurons into projection neurons (pyramidal and spiny stellate) and interneurons (non-pyramidal, local circuit neurons).     

Ultrastructure of visual callosal neurons in cat identified by retrograde axonal transport of horseradish peroxidase

Summary The ultrastructure of neurons at the border of areas 17 and 18 of the visual cortex of the cat was studied by the combined use of the retrograde transport of horseradish peroxidase (HRP) and electron microscopy. Callosal neurons were retrogradely labelled by injecting HRP at the 17/18 border region of the contralateral hemisphere. They were found mainly in layer III but also in IV and VI. They were most commonly pyramidal cells and less often large, spiny stellate cells. Pyramidal callosal neurons received only symmetrical synapses on their soma and mainly symmetrical (but a few asymmetrical) synapses on their dendritic shafts. Their abundant spines received asymmetrical synapses. The stellate cells were contacted by moderate numbers of symmetrical and asymmetrical axodendritic and axosomatic synapses and also had asymmetrical axospinous contacts. We propose that the callosal stellate neurons consist of a class of large spiny stellates, recognizable by light and electron microscopic criteria.The work described in this paper forms part of a study for a doctoral dissertation in the University of Lausanne by J. P. Hornung.     

Morphological correlates of visual field transformation in the corpus callosum

Horseradish peroxidase (HRP) injections at the boundary of areas 17 and 18 in cat visual cortex resulted in retrograde labelling of neurones located in contralateral areas 17 and 18. The HRP-positive neurones were spread over a region located around the representation of the vertical meridian; they were most numerous in the locus for area centralis. Layer III pyramidal cells and stellate cells in upper layer IV were identified as the principal neurones of origin of the visual callosal pathway.     

The spiny stellate neurons in layer IV of the human auditory cortex. A Golgi study

The spiny stellate neurons have been studied by the Golgi method in the auditory koniocortex and parakoniocortex of man. Spiny stellate cells are a consistent though not very common component of layer IV. They are not confined to specific sublayers but occur at all depths of layer IV, and also in layer IIIc. Spiny stellate cells in the auditory areas show a great variety of their dendritic arborization pattern. The presence of all intermediate forms between small pyramidal cells--which constitute the dominant cell type in layer IV and which display an extraordinary heteromorphism--and spiny stellate cells shows the close kinship between both neuronal types. The morphology and distribution of spines along the dendrites of spiny stellate neurons are similar to those of the small pyramidal cells of the same layer. The axons, which were impregnated only in their proximal portions, mostly descend, giving rise to recurrent ascending collaterals, but initially ascending axons do also occur. Spiny stellate neurons are present in the different cytoarchitectonic areas examined, and thus they are not confined to the auditory koniocortex.     

Local circuits targeting parvalbumin-containing interneurons in layer IV of rat barrel cortex

Interactions between inhibitory interneurons and excitatory spiny neurons and also other inhibitory cells represent fundamental network properties which cause the so-called thalamo-cortical response transformation and account for the well-known receptive field differences of cortical layer IV versus thalamic neurons. We investigated the currently largely unknown morphological basis of these interactions utilizing acute slice preparations of barrel cortex in P19-21 rats. Layer IV spiny (spiny stellate, star pyramidal and pyramidal) neurons or inhibitory (basket and bitufted) interneurons were electrophysiologically characterized and intracellularly biocytin-labeled. In the same slice, we stained parvalbumin-immunoreactive (PV-ir) interneurons as putative target cells after which the tissue was subjected to confocal image acquisition. Parallel experiments confirmed the existence of synaptic contacts in these types of connection by correlated light and electron microscopy. The axons of the filled neurons differentially targeted barrel PV-ir interneurons: (1) The relative number of all contacted PV-ir cells within the axonal sphere was 5–17% for spiny (n = 10), 32 and 58% for basket (n = 2) and 12 and 13% for bitufted (n = 2) cells. (2) The preferential subcellular site which was contacted on PV-ir target cells was somatic for four and dendritic for five spiny cells; for basket cells, there was a somatic and for bitufted cells a dendritic preference in each examined case. (3) The highest number of contacts on a single PV-ir cell was 9 (4 somatic and 5 dendritic) for spiny neurons, 15 (10 somatic and 5 dendritic) for basket cells and 4 (1 somatic and 3 dendritic) for bitufted cells. These patterns suggest a cell type-dependent communication within layer IV microcircuits in which PV-ir interneurons provide not only feed-forward but also feedback inhibition thus triggering the thalamo-cortical response transformation.     

Morphology of visual callosal neurons with different locations,contralateral targets or patterns of development

In kittens, callosally projecting neurons were labeled by retrograde transport of FITC- (fluorescein isothiocyanate)- and TRITC- (tetramethylrhodamine isothiocyanate)-conjugated latex microspheres injected in two different visual areas (17, 17/18, 19, or postero-medial lateral suprasylvian; PMLS) at postnatal day 3. At postnatal day 57 more than 1200 labeled neurons in visual cortical areas were intracellularly injected with 3% lucifer yellow (LY) in perfusion-fixed slices of the contralateral hemisphere. The distribution of labeled neurons was charted, and LY-filled neurons were classified on the basis of their area and layer of location, and dendritic pattern. The dendritic arbors of 120 neurons were computer reconstructed. For the basal dendrites of supragranular pyramidal neurons a statistical analysis of number of nodes, internodal and terminal segment lengths, and total dendritic length was run relative to the area of location and axonal projection. Connections were stronger between homotopic than between heterotopic areas. Overall tangential and laminar distributions depended on the area injected. Qualitative morphological differences were found among callosally projecting neurons, related to the area of location, not to that of projection. In all projections from areas 17 and 18, pyramidal and spinous stellate neurons were found in supragranular layers. In contrast, spinous stellate neurons lacked in projections from area 19, 21a, PMLS and postero-lateral lateral suprasylvian (PLLS). In all areas, the infragranular neurons showed heterogeneous typology, but in PMLS no fusiform cells were found. Quantitative analysis of basal dendrites did not reveal significant differences in total dendritic length, terminal, or intermediate segment length among neurons in area 17 or 18, and this was related to whether they projected to contralateral areas 17–18 or PMLS. All injections produced exuberant labeling in area 17. No differences could be found between neurons in area 17 (with transient axons through the corpus callosum) and neurons near the 17/18 border (which maintain projections to the corpus callosum). In conclusion, morphology of callosally projecting neurons seems to relate more to intrinsic specificities in the cellular composition of each area than to the area of contralateral axonal projection or the fate of callosal axons.     

Morphology of spiny neurons in the human entorhinal cortex: intracellular filling with lucifer yellow

The present study was designed to investigate the morphology of spiny neurons in the human entorhinal cortex. Coronal entorhinal slices (n = 67; 200 microm thick) were obtained from autopsies of three subjects. Spiny neurons (n = 132) filled with Lucifer Yellow were analysed in different subfields and layers of the entorhinal cortex. Based on the shape of the somata and primary dendritic trees, spiny neurons were divided into four morphological categories; (i) classical pyramidal, (ii) stellate, (iii) modified stellate, and (iv) horizontal tripolar cells. The morphology of filled neurons varied more in different layers than in the different subfields of the entorhinal cortex. In layer II, the majority (81%) of spiny neurons had stellate or modified stellate morphology, but in the rostromedial subfields (olfactory subfield and rostral subfield) there were also horizontal tripolar neurons. Dendritic branches of layer II neurons extended to layer I (94%) and to layer III (83%). Unlike in layer II, most (74%) of the filled neurons in layers III, V and VI were classical pyramidal cells. The majority of pyramidal cells in the superficial portion of layer III had dendrites that extended up to layer II, occupying the space between the neuronal clusters. Some dendrites reached down to the deep portion of layer III. Apical dendrites of layer V and VI pyramidal cells traveled up to the deep portion of layer III.Our data indicate that the morphology of spiny neurons in different layers of the human entorhinal cortex is variable. Vertical extension of dendritic branches to adjacent layers supports the idea that inputs terminating in a specific lamina influence target cells located in various entorhinal layers. There appears to be more overlap in the dendritic fields between superficial layers II and III than between the superficial (II/III) and deep (V/VI) layers, thus supporting the idea of segregation of information flow targeted to the superficial or deep layers in the human entorhinal cortex.     

Synaptic relationships of a type of GABA-immunoreactive neuron (clutch cell), spiny stellate cells and lateral geniculate nucleus afferents in layer IVC of the monkey striate cortex

The precise stimulus specificity of striate cortical neurons is strongly influenced by processes involving gamma-aminobutyric acid (GABA). In the visual cortex of the monkey most afferents from the lateral geniculate nucleus terminate in layer IVC. We identified a type of smooth dendritic neuron (clutch cell) that was immunoreactive for GABA, and whose Golgi-impregnated dendrites and axon were largely restricted to layer IVC beta. The slightly ovoid somata were 8-12 micron by 12-15 micron and the dendritic field was often elongated, extending 80-200 micron in one dimension. The axonal field was 100-150 micron in diameter and densely packed with large bulbous boutons. Although mainly located in IVC beta both the dendritic and axonal processes entered IVC alpha. Fine structural features of GABA-immunoreactive and-impregnated clutch cells and impregnated spiny stellate cells were compared. Clutch cells had more cytoplasm, more densely packed mitochondria and endoplasmic reticulum, and made type II as opposed to type I synapses. A random sample of 159 elements postsynaptic to three clutch cells showed that they mainly terminated on dendritic shafts (43.8-58.5%) and spines (20.8-46.3%), rather than somata (10-17%). The majority of the postsynaptic targets were characteristic of spiny stellate cells. This was directly demonstrated by studying synaptic contacts between an identified GABA positive clutch cell and the dendrites and soma of an identified spiny stellate cell. The termination of clutch cells mainly on dendrites and spines of spiny stellate cells suggests that they interact with other inputs to the same cells. Following an electrolytic lesion in the ipsilateral lateral geniculate nucleus we examined the distribution of degenerating terminals on three identified spiny stellate neurons in layer IVC beta. Out of eight synapses from the lateral geniculate nucleus one was on a dendritic shaft, the rest on spines. Only a small fraction of all synapses on the cells were from degenerating boutons. A clutch cell within the area of dense terminal degeneration was not contacted by terminals from the lateral geniculate nucleus. The results show that layer IVC in the monkey has a specialized GABAergic neuron that terminates on spiny stellate cells monosynaptically innervated from the lateral geniculate nucleus. Possible functions of clutch cells may include inhibitory gating of geniculate input to cortex; maintenance of the antagonistic subregions in the receptive fields; and the creation from single opponent of double colour opponent receptive fields.     

The thalamic projection to cat visual cortex: ultrastructure of neurons identified by Golgi impregnation or retrograde horseradish peroxidase transport

Specific afferents to the primary visual cortex of the cat were identified by electron microscopy after electrolytic lesions of the lateral geniculate nucleus. Postsynaptic target neurons were marked by either Golgi impregnation or horseradish peroxidase retrogradely transported from the opposite visual cortex. The type and the location of identified neurons were determined by light microscopy before further investigation by electron microscopy. Different pyramidal neurons have relatively homogeneous ultrastructural characteristics, but non-pyramidal neurons, divisible into large and small, spiny, sparsely-spiny and non-spiny, vary in their synaptology. Thalamo-cortical afferents have been found to all neuronal types examined in layer IV and lower layer III, including horseradish peroxidase-filled callosal cells. These include pyramidal cells of layer III which receive thalamo-cortical afferents mainly on spines of apical and basal dendrites, but also on basal dendritic shafts. Layer V pyramids receive thalamo-cortical input to apical dendritic spines. All types of layer IV non-pyramidal neurons studied are contacted by geniculate terminals on dendritic spines and shafts and directly on the cell body.We conclude that input to the visual cortex is organized on both hierarchical and parallel bases.     

The projection of the lateral geniculate nucleus to area 17 of the rat cerebral cortex. V. Degenerating axon terminals synapsing with Golgi impregnated neurons

Summary The sites of termination of afferents from the lateral geniculate nucleus to layer IV and lower layer III in area 17 of the rat visual cortex have been determined by use of a combined degeneration—Golgi/EM technique. Degeneration of geniculocortical axon terminals was produced by making lesions in the lateral geniculate body. After the animals had been allowed to survive for two days, the ipsilateral visual cortex was removed and impregnated by the Golgi technique. Suitably impregnated neurons and their processes in layer IV and lower layer III were then gold-toned and deimpregnated for examination in the electron microscope. A search was made for synapses between degenerating axon terminals and the gold-labelled postsynaptic neurons.Geniculocortical synapses were found to involve: (1) the spines of basal dendrites, as well as those of proximal shafts and collaterals of apical dendrites of layer III pyramidal neurons; (2) the spines of the apical dendritic shafts and collaterals of layer V pyramidal neurons; (3) the perikaryon and dendritic spines of a sparsely-spined stellate cell; and (4) the perikaryon and dendrites of a smooth, bitufted stellate cell. In view of this variety of postsynaptic elements it is suggested that all parts of the perikarya and dendrites of neurons contained in layer IV and lower layer III which are capable of forming asymmetric synapses can be postsynaptic to the thalamic input.Finally, an analysis of the known neuronal interrelations within the rat visual cortex is presented.     

A Golgi deimpregnation study of neurons in the rhesus monkey visual cortex (Areas 17 and 18)

Summary The morhological features of 298 neurons impregnated according to Golgi-Kopsch in areas 17 and 18 of Macaca mulatta were analyzed, and the same neurons were deimpregnated to visualize structural details of the somata in different types of neurons. The following cell types were investigated: Pyramidal and pyramid-like cells, spiny stellate cells, double bouquet cells, bipolar cells, chandelier cells, neurogliaform cells, basket and related cells. This procedure allows the evaluation of the nuclear-cytoplasmic proportion and the position of the nucleus besides shape and size of the cell body. Pyramidal and pyramid-like cells (N=43), spiny stellate cells (N=26), basket and related cells (N=126) are variable in these features. A positive correlation between soma size and width of the cytoplasm is found in pyramidal, pyramid-like cells and spiny stellate cells. With the exception of some large somata in both these types of neurons the nucleus is found in a central position. Double bouquet cells (N=6), bipolar cells (N=13) and chandelier cells (N=11) exhibit small cytoplasmic rims and centrally located nuclei. The small somata of neurogliaform cells (N=37), however, and the small to very large somata of basket and related cells show broad cytoplasmic portions surrounding the eccentrically located nuclei. These findings allow the identification of different neuronal types in Nisslstained sections on the basis of these soma features. This is a prerequisite for further detailed quantitative studies on the laminar distribution of different neuronal types in the visual cortex of the monkey.     

Corticocortical connections to the motor cortex from the posterior parietal lobe (areas 5a, 5b, 7) in the cat demonstrated by the retrograde axonal transport of horseradish peroxidase

Summary Neurons in the parietal region of the cerebral cortex, projecting to the ipsilateral distal forelimb area of the motor cortex (area 4) were identified in the cat brain using the horseradish peroxidase (HRP) retrograde tracing method. After making microinjections of HRP into the distal forelimb area of the motor cortex, clusters of HRP-labeled cell bodies were observed in different regions of the ipsilateral parietal cortex. In particular these clusters of labeled cells were found in areas 5a, 5b and 7. The area 5a cluster is formed from closely packed irregularly-shaped cells, the area 5b cluster is made up of dispersed medium-sized pyramidal cells, while area 7 contains a cluster of widely dispersed small pyramidal cells. Typically, labeled cell bodies were found in lamina III of cortex. Labeled cell bodies were neither observed in the contralateral cortex nor in the visual cortex (areas 17, 18 and 19). Since parietal cortex receives projections from primary somatosensory and visual cortex, the projections from parietal to motor cortex may well form the neural substrate for the processing of convergent sensory information used in voluntary movements.     

Morphological evidence for callosally projecting nonpyramidal neurons in rat visual cortex

Summary This investigation shows that some of the callosally projecting neurons in rat visual cortex are nonpyramidal cells. Callosally projecting neurons were labeled by injections of horseradish peroxidase (HRP) into the area 17/18 a border zone of the contralateral hemisphere. The retrogradely transported HRP was visualized with diaminobenzidine or with tetramethylbenzidine. In some of the labeled neurons the reaction product was diffuse, so that the neurons had a Golgi-like appearance, but in others the reaction product was granular, or punctate. The majority of neurons with a Golgi-like appearance were pyramidal cells, but one callosally projecting neuron from layer V area 18 a was confirmed by electron microscopy to be a nonpyramidal neuron. This dearth of well-filled nonpyramidal cells suggested that callosally projecting nonpyramidal neurons may not transport sufficient HRP to show Golgi-like filling, and so punctately labeled neurons from areas 17, 18 a and 18 b were examined. Reacted sections from areas 17, 18 a and 18 b of control animals, into which no tracer had been injected, were also examined, but in these control preparations no granules similar to the HRP granules within the neuronal profiles of the experimental animals were encountered. In methylene blue-stained 1-m sections, neuronal profiles from the control animals possessed only blue staining lysosomes, while neuronal profiles from the experimental animals exhibited both lysosomes and HRP granules. It was determined, from the counts of HRP granules in neurons from the experimental animals, that in selected regions of areas 17, 18 a, and 18 b similar percentages of the pyramidal and nonpyramidal neuronal populations (ranging from 100% to 34%) contained HRP granules, and so had callosally projecting axons. However, most callosally projecting nonpyramidal neurons had far fewer HRP granules than the pyramidal neurons, again indicating that they transport less HRP. This could account for the fact that callosally projecting nonpyramidal neurons only rarely show a Golgi-like filling, and this could be one reason why such cells have been overlooked in most previous studies.     

The anatomical substrate of callosal messages from SI and SII in the cat

Summary 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.R. C. performed part of the work while on leave at the Institute of Anatomy, University of Lausanne.     

Neuromodulatory role of acetylcholine in visually-induced cortical activation: behavioral and neuroanatomical correlates

Acetylcholine is released in the primary visual cortex during visual stimulation and may have a neuromodulatory role in visual processing. The present study uses both behavioral and functional neuroanatomy investigations to examine this role in the rat. In the first set of experiments the cholinergic system was lesioned with 192 immunoglobulin G (IgG) saporin and the visual acuity and performance in a visual water maze task were assessed. The cholinergic lesion did not affect the visual acuity measured pre- and post-lesion but it did reduce the efficiency to learn a novel orientation discrimination task measured post-lesion. In order to better understand the involvement of the cholinergic system in the neuronal activity in the visual cortex c-Fos expression induced by patterned visual stimulation was further investigated. Results obtained following lesion of the cholinergic fibers (192 IgG-saporin or quisqualic acid), muscarinic inhibition (scopolamine), or NMDA receptor inhibition (CPP) were compared with control conditions. Double and triple immunolabeling was used in order to determine the neurochemical nature of the activated cortical cells. The results demonstrated that patterned stimulation elicited a significant increase in c-Fos immunolabeled neurons in layer IV of the contralateral primary visual cortex to the stimulated eye which was completely abolished by cholinergic fibers lesion as well as scopolamine administration. This effect was independent of NMDA receptor transmission. The c-Fos activation was predominantly observed in the glutamatergic spiny stellate cells and less frequently in GABAergic interneurons. Altogether, these results demonstrate a strong involvement of the basal forebrain cholinergic system in the modulation of post-synaptic visual processing, which could be related to cognitive enhancement or attention during visual learning.     

Turtle hippocampal cortex contains distinct cell types, burst-firing neurons, and an epileptogenic subfield

The dorsal and medial telencephalon of reptiles consists of a simple trilaminar cortex. The turtle dorsal cortex has been identified as a favorable physiological preparation that may bear a phylogenetic relationship to mammalian neocortex. While anatomical studies have likened the reptilian medial cortical region to mammalian hippocampus, its physiological properties have not been explored. We therefore used intracellular and extracellular recording techniques to examine the cellular and synaptic physiology of turtle "hippocampal" or medial cortex. Turtle medial cortex contains two principal classes of neurons, pyramidal cells and stellate neurons. Recordings with Lucifer yellow CH (LY)-filled microelectrodes allowed us to correlate the physiological properties of medial cortical neurons with their cellular morphology. Pyramidal neurons were situated in a single cellular layer and had spiny apical dendrites extending into the molecular layer. These cells fired relatively long-duration action potentials (APs) and showed frequency adaptation to suprathreshold current pulse injections. Stellate cells were usually found in the subcellular and molecular layers and had aspiny dendrites. In contrast to pyramidal cells, they fired brief APs and displayed no frequency adaptation. A discrete population of cells in the dorsal portion of medial cortex (DMC) was capable of bursting endogenously or in response to synaptic activation. Bursts usually contained an underlying slow depolarization and often occurred at regular intervals. Intracellular LY injections confirmed that these cells were pyramidal in morphology. Electrical stimulation of afferent fibers revealed that pyramidal cells and stellate neurons differed in their synaptic responses. In ventral medial cortex (VMC), afferent stimulation evoked a multiphasic response in most pyramidal cells, whereas stellate cells were synaptically excited. Orthodromic activation of DMC bursting cells resulted in a powerful excitation--often a short burst--and subsequent inhibition. Stellate neurons in DMC also had a biphasic synaptic response consisting of both an early excitation and a late inhibition. Experiments using intracellular chloride (Cl-) injection or focal bicuculline application suggested that part of the inhibitory component of the pyramidal cell synaptic response was dependent on a gamma-aminobutyric acid (GABA)-mediated increase in Cl- conductance. These results correlated with our immunohistochemical studies that revealed the presence of GABAergic neurons in medial cortex.(ABSTRACT TRUNCATED AT 400 WORDS)     

Morphology of neurons in the white matter of the adult human neocortex

Summary Neurons in the human cerebral cortical white matter below motor, visual, auditory and prefrontal orbital areas have been studied with the Golgi method, immunohistochemistry and diaphorase histochemistry. The majority of white matter neurons are pyramidal cells displaying the typical polarized, spiny dendritic system. The morphological variety includes stellate forms as well as bipolar pyramidal cells, and the expression of a certain morphological phenotype seems to depend on the position of the neuron. Spineless nonpyramidal neurons with multipolar to bitufted dendritic fields constitute less than 10% of the nuerons stained for microtubule associated protein (MAP-2). Only 3% of the MAP-2 immunoreactive neurons display nicotine adenine dinucleotide-diaphorase activity. The white matter pyramidal neurons are arranged in radial rows continuous with the columns of layer VI neurons. Neuron density is highest below layer VI, and decreases with increasing distance from the gray matter. White matter neurons are especially abundant below the primary motor cortex, and are least frequent below the visual cortex area 17. In contrast to other mammalian species, the white matter neurons in man are not only present during development, but persist throughout life.