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.     

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.     

Number estimates of neuronal phenotypes in layer II of the medial entorhinal cortex of rat and mouse

Modelling entorhinal function or evaluating the consequences of neuronal losses which accompany neurodegenerative disorders requires detailed information on the quantitative cellular composition of the normal entorhinal cortex. Using design-based stereological methods, we estimated the numbers, proportions, densities and sectional areas of layer II cells in the medial entorhinal area (MEA), and its constituent caudal entorhinal (CE) and medial entorhinal (ME) fields, in the rat and mouse. We estimated layer II of the MEA to contain ∼58,000 neurons in the rat and ∼24,000 neurons in the mouse. Field CE accounted for more than three-quarters of the total neuron population in both species. In the rat, layer II of the MEA is comprised of 38% ovoid stellate cells, 29% polygonal stellate cells and 17% pyramidal cells. The remainder is comprised of much smaller populations of horizontal bipolar, tripolar, oblique pyramidal and small round cells. In the mouse, MEA layer II is comprised of 52% ovoid stellate cells, 22% polygonal stellate cells and 14% pyramidal cells. Significant species differences in the proportions of ovoid and polygonal stellate cells suggest differences in physiological and functional properties. The majority of MEA layer II cells contribute to the entorhinal-hippocampal pathways. The degree of divergence from MEA layer II cells to the dentate granule cells was similar in the rat and mouse. In both rat and mouse, the only dorsoventral difference we observed is a gradient in polygonal stellate cell sectional area, which may relate to the dorsoventral increase in the size and spacing of individual neuronal firing fields. In summary, we found species-specific cellular compositions of MEA layer II, while, within a species, quantitative parameters other than cell size are stable along the dorsoventral and mediolateral axis of the MEA.     

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.     

Electrophysiological classes of cat primary visual cortical neurons in vivo as revealed by quantitative analyses

To facilitate the characterization of cortical neuronal function, the responses of cells in cat area 17 to intracellular injection of current pulses were quantitatively analyzed. A variety of response variables were used to separate the cells into subtypes using cluster analysis. Four main classes of neurons could be clearly distinguished: regular spiking (RS), fast spiking (FS), intrinsic bursting (IB), and chattering (CH). Each of these contained significant subclasses. RS neurons were characterized by trains of action potentials that exhibited spike frequency adaptation. Morphologically, these cells were spiny stellate cells in layer 4 and pyramidal cells in layers 2, 3, 5, and 6. FS neurons had short-duration action potentials (<0.5 ms at half height), little or no spike frequency adaptation, and a steep relationship between injected current intensity and spike discharge frequency. Morphologically, these cells were sparsely spiny or aspiny nonpyramidal cells. IB neurons typically generated a low frequency (<425 Hz) burst of spikes at the beginning of a depolarizing current pulse followed by a tonic train of action potentials for the remainder of the pulse. These cells were observed in all cortical layers, but were most abundant in layer 5. Finally, CH neurons generated repetitive, high-frequency (350-700 Hz) bursts of short-duration (<0.55 ms) action potentials. Morphologically, these cells were layer 2-4 (mainly layer 3) pyramidal or spiny stellate neurons. These results indicate that firing properties do not form a continuum and that cortical neurons are members of distinct electrophysiological classes and subclasses.     

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.     

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)     

Acute stress-induced neuronal plasticity in the corticoid complex of 15-day-old chick,Gallus domesticus

Several studies conducted on chicken have shown that a single stress exposure may impair or improve memory as well as learning processes. However, to date, stress effects on neuronal morphology are poorly investigated wherefore it was of interest to evaluate this further in chicks. Thus, the present study aims to investigate the role of single acute stress (AS) of 24 h food and water deprivation in neuronal plasticity in terms of spine density of the corticoid complex (CC) in 15-day-old chick, Gallus domesticus, by using three neurohistological techniques: Cresyl Violet, Golgi Colonnier, and Golgi Cox technique. The dorsolateral surface of the cerebral hemisphere is occupied by CC which can be differentiated into two subfields: an intermediate corticoid (CI) subfield (arranged in layers) and a dorsolateral corticoid (CDL) subfield. Based on different criteria such as soma shape, dendritic branching pattern, and dendritic spine density, two main moderately spinous groups of neuronal cells were observed in the CC, namely, projection neurons (comprising of multipolar and pyramidal neurons) and stellate neurons. In the present study, the stellate neurons have shown a significant decrease as well as an increase in their spine density in both CI and CDL subfields, whereas the multipolar neurons had shown a significant increase in their spine density in the CDL region only. The present study shows that AS induces neuronal plasticity in terms of spine density in both CI and CDL neurons. The morphological changes in the form of decreased dendritic branches due to stress have been observed in the CI region in comparison to CDL region, which could be linked to more effect of stress in this region. The avian CDL corresponds to the entorhinal cortex of mammals on the basis of neuronal morphology and bidirectional connections between adjacent areas. The projection neurons increase their branches and also their spine number to cope with the stress effects, while the stellate neurons show contrasting effect in their spine density. Therefore, this study will establish that slight modifications in natural stimuli or environmental changes faced by the animal may affect their dorsolateral forebrain which shows neuronal plasticity that help in the development of an adaptive capacity of the animal to survive under changing environmental conditions.     

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.     

Somatostatin and vasoactive intestinal polypeptide-like immunoreactive cells and terminals in the retrohippocampal region of the rat brain

The retrohippocampal region of the rat brain was analyzed by using immunohistochemistry with specific antibodies against somatostatin (SOM) and vasoactive intestinal polypeptide (VIP). Specifically immunoreactive neurons and terminal processes were labeled with either the anti-SOM or anti-VIP antiserum and they were referred to as SOM-like immunoreactive (SOM-LI) or VIP-like immunoreactive (VIP-LI) neurons and processes, respectively. The retrohippocampal region was rich in neuronal cell bodies and terminal processes showing immunoreactivity for SOM and VIP. In the entorhinal area SOM-LI neurons were located mainly in layers IV through VI and the VIP-LI neurons were found mainly in layers I through III. Thick (70-120 microns) sections treated with the immunoperoxidase method to achieve a Golgi-like staining pattern showed that cytological differences existed between SOM- and VIP-positive neurons. SOM-LI neurons were usually multipolar, fusiform, or occasionally pyramidal while VIP-LI neurons were usually bipolar, stellate, or fusiform. SOM-LI and VIP-LI axons and preterminal processes were differentially distributed within the laminae of the retrohippocampal region. VIP-LI terminals were found throughout all layers except layer I. SOM-LI terminals were found primarily in the molecular layers of all areas, layer IV of the medical and lateral entorhinal areas, and in the angular bundle. Thus, SOM-LI and VIP-LI neurons are distinguished by their morphology and their different distribution within the cortical layers and areas of the retrohippocampal region.     

Somatostatin immunoreactive neurons in rat visual cortex: A light and electron microscopic study

Summary Somatostatin immunoreactive neurons in rat visual cortex were examined in the light and electron microscopes using an antibody to the tetradecapeptide form of somatostatin. Somatostatin immunoreactive neurons were found to belong only to non-pyramidal classes. They are of five main types: multipolar neurons with either thin or thick dendrites; small and large bipolar neurons; bitufted neurons; horizontal neurons; and neurons in the subcortical white matter. Of the immunoreactive neurons, multipolar neurons are the most common and account for 30% of the population, while bipolar and bitufted neurons make up 25% and 15% of the immunoreactive population, respectively; the least common somatostatin immunoreactive neurons are the horizontal and subcortical white matter neurons. Occasional multipolar neurons with thick dendrites have a prominent ascending dendrite so that they resemble pyramidal cells in the light microscope, but electron microscopic examination confirms that, like all other somatostatin-positive cells, they are non-pyramidal neurons, for they have both symmetric and asymmetric synapses on their cell bodies.Somatostatin-positive neurons are distributed among all the cortical layers and the subcortical white matter but they are more common in two laminae, one coinciding with layer II/III and the other with layers V and VI. The multipolar and bipolar neurons are distributed in similar proportions in these upper and lower cortical laminae, while bitufted neurons are more common in upper laminae and horizontal neurons are predominantly located in layer VI.     

Microstructure of the neocortex: comparative aspects

The appearance of the neocortex, its expansion, and its differentiation in mammals, represents one of the principal episodes in the evolution of the vertebrate brain. One of the fundamental questions in neuroscience is what is special about the neocortex of humans and how does it differ from that of other species? It is clear that distinct cortical areas show important differences within both the same and different species, and this has led to some researchers emphasizing the similarities whereas others focus on the differences. In general, despite of the large number of different elements that contribute to neocortical circuits, it is thought that neocortical neurons are organized into multiple, small repeating microcircuits, based around pyramidal cells and their input-output connections. These inputs originate from extrinsic afferent systems, excitatory glutamatergic spiny cells (which include other pyramidal cells and spiny stellate cells), and inhibitory GABAergic interneurons. The problem is that the neuronal elements that make up the basic microcircuit are differentiated into subtypes, some of which are lacking or highly modified in different cortical areas or species. Furthermore, the number of neurons contained in a discrete vertical cylinder of cortical tissue varies across species. Additionally, it has been shown that the neuropil in different cortical areas of the human, rat and mouse has a characteristic layer specific synaptology. These variations most likely reflect functional differences in the specific cortical circuits. The laminar specific similarities between cortical areas and between species, with respect to the percentage, length and density of excitatory and inhibitory synapses, and to the number of synapses per neuron, might be considered as the basic cortical building bricks. In turn, the differences probably indicate the evolutionary adaptation of excitatory and inhibitory circuits to particular functions.     

Neurons in the ventral subiculum, amygdala and entorhinal cortex which project to the nucleus accumbens: Their input from somatostatin-immunoreactive boutons

Neurons in the hippocampus, amygdala and entorhinal cortex which project to the nucleus accumbens were labelled retrogradely following injection of horseradish peroxidase. The injections were targetted on the medial part of the nucleus accumbens, but some injection sites included the whole nucleus. Projection neurons in all three areas were found to be spiny, and from the entorhinal cortex and ventral subiculum of the hippocampus they were pyramidal neurons.Somatostatin (S28_1–12)-immunoreactive neurons were found in all parts of the three limbic areas examined. They were found to have various morphologies, but in the electron microscope all had the ultrastructural characteristics of interneurons. In the hippocampus the stratum lacunosum was found to contain the most immunoreactive fibres while most cells lay in the stratum oriens. In the amygdala the densest staining for both cells and fibres was in the central nucleus. In the entorhinal cortex somatostatin-immunoreactive fibres and cells seemed to have no preferential distribution.Examination of somatostatin-immunoreactive profiles in the electron microscope revealed that the majority of synaptic contacts were made with dendrites, many of which were spine-bearing.In the light microscope somatostatin-immunoreactive fibres could be seen to lie near the somata and proximal dendrites of neurons that projected to the nucleus accumbens. In the electron microscope it was found that somatostatin-immunoreactive boutons were in symmetrical synaptic contact with the somata and proximal dendrites of neurons in the ventral subiculum, entorhinal cortex and amygdala which project to the nucleus accumbens.     

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.     

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.     

Development of full-length Trk B-immunoreactive structures in the hippocampal formation of the macaque monkey

Distribution and morphological changes of cells containing the signal transducing neurotrophin receptor, full-length Trk B (fl-Trk B), were investigated in the hippocampal formation of the macaque monkey between embryonic day 140 and the adult stage. Western blot analysis showed that one main protein band, which migrated at 141 kDa, was detected in both the embryonic and adult hippocampal formation. In the pyramidal cells in CA1 and CA3 subfields, the subiculum, and the entorhinal cortex, fl-Trk B-immunoreactive dendrites were observable in the embryonic stage. In contrast, in the granule cells of the dentate gyrus, few dendrites were immunoreactive during embryonic and early developmental stages. This difference may be due to the later growth of the granule cells of the dentate gyrus. The existence of fl-Trk B immunoreactivity in the cell body and dendrites in the embryonic hippocampal neurons, suggests that BDNF and/or NT4/5 act on the hippocampal cells by autocrine/paracrine mechanisms. In the entorhinal cortex, fl-Trk B immunoreactivity became localized in the stellate cells in layer II and the pyramidal cells in layers III, V and VI in adulthood. This indicates that BDNF and/or NT4/5 are important for the maintenance of the projection neurons in the entorhinal cortex at the adult stage. The strongest fl-Trk B immunoreactivity in the hippocampal neurons occurred at postnatal month 4, corresponding to the period of greatest synapse production in the monkey hippocampus, suggesting that BDNF and/or NT4/5 with fl-Trk B may play a role in synapse formation in the monkey hippocampus.     

Membrane properties of identified lateral and medial perforant pathway projection neurons

The physiological characteristics of neurons that project to the hippocampus and dentate gyrus via the medial perforant pathway (projection neurons) are well known, but the characteristics of neurons that project to these areas via the lateral perforant pathway (projection neurons) are less well known. We have used retrograde tracing and whole-cell recording in brain slices to compare the membrane and firing properties of medial perforant pathway and lateral perforant pathway projection neurons in layer II of the medial and lateral entorhinal cortex. The properties of medial perforant pathway projection neurons were identical to those reported previously for spiny stellate neurons in the medial entorhinal cortex. In contrast, lateral perforant pathway projection neurons were characterized by a higher input resistance, a lack of time-dependent inward (anomalous) rectification, and a lack of prominent depolarizing spike afterpotentials. Voltage-clamp recordings suggest that the absence of anomalous rectification in lateral perforant pathway projection neurons is due to smaller hyperpolarization activated cation currents in these cells, and the lack of depolarizing afterpotential may be due to smaller low-threshold calcium currents. Persistent sodium current was also smaller in lateral perforant pathway projection neurons, but the difference in persistent sodium current between medial perforant pathway and lateral perforant projection neurons was much less pronounced than the difference in low voltage activated currents. These results underscore the functional differences between the medial entorhinal cortex and lateral entorhinal cortex, and may help to explain the differing abilities of these cortical areas to participate in certain types of network activity.     

The callosal projection in cat visual cortex as revealed by a combination of retrograde tracing and intracellular injection

Summary The neuronal composition of callosally projecting cells in cat visual cortex was determined with a combination of retrograde labelling and intracellular injection. Fluorescent tracers were stereotaxically injected into the proximity of the area 17/18 border, corresponding to the representation of the visual vertical meridian. In fixed slice preparations of homotopic regions of the contralateral hemisphere retrogradely labelled cells were filled with Lucifer Yellow. Of more than a hundred injected cells a morphological variety of pyramidal cells, located in cortical layers II–IV and VI, constituted the prevalent cell class in the contralateral projection. A minor proportion of spiny stellate cells was encountered in layer IV. Despite the presence of a contralaterally projecting smooth stellate cell, presumed to be a basket cell, it is concluded that the efferents to contralateral visual cortex predominantly arise from pyramidal and spiny stellate cells. Thus, in agreement with findings from anterograde degeneration studies, the interhemispheric pathway most likely conveys a direct excitatory input to postsynaptic target cells.