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PubMed1996

Temporal and spatial appearance of the membrane cytoskeleton and perineuronal nets in the rat neocortex.

Parvalbumin-immunoreactive interneurons are surrounded by perineuronal nets, containing molecules of the extracellular matrix (e.g. tenascin-R). Furthermore, they seem to have a special cytoskeleton composed of, among others, ankyrinR and beta Rspectrin. In the present developmental study we showed that the intracellular markers parvalbumin, ankyrinR and beta Rspectrin as well as Vicia Villosa agglutinin, an extracellular marker for perineuronal nets, appeared in the second postnatal week. In the third postnatal week, ankyrinR and beta R spectrin were present in the parvalbumin-positive interneurons. Tenascin-R appeared in a similar topographic distribution as the intracellular markers. The adult pattern was established upon the end of the fourth postnatal week. Our results indicate that cytoskeletal maturity maybe a prerequisite for the organization of perineuronal nets of extracellular matrix.

Animals↗
PubMed1996

Antibodies to glutamate, aspartate and glycyl-D-aspartate reversibly suppress stimulus-evoked, extracellularly recorded responses in slices of rat neocortex.

Polyclonal antibodies raised against glutamate, aspartate and the dipeptide, glycyl-D-aspartate were dissolved in artificial cerebrospinal fluid (aCSF) and administered at concentrations as low as 0.05% to slices of prefrontal cortex maintained in vitro. These antisera caused a reversible attenuation of evoked field potentials and/or single-unit activity recorded extracellularly following the delivery of shocks to the underlying white matter, or to cortical layer IV. To the best of our knowledge, this result provides the first demonstration using electrophysiological recording of the use of a transmitter-specific antibody as a blocker of synaptic transmission in living slices of the central nervous system (CNS). The results lend support to the suggestion that glutamate, aspartate, and a molecule related closely to glycyl-D-aspartate, are involved in synaptic transmission at major pathways within prefrontal cortex.

Animals↗
PubMed1982

Excitatory effect of acetylcholine on different types of neurons in the first somatosensory neocortex of the rat: laminar distribution and pharmacological characteristics.

In rats anaesthetized with either urethane, pentobarbital or fluothane the effects of acetylcholine, cholinergic agonists and antagonists (applied by iontophoresis) were studied on single cortical neurons of first somatosensory region. The laminar distribution of the neurons excited by acetylcholine was determined by the reconstruction of each electrode track based on a dye-deposit made at the last recording site. Neurons were identified using antidromic stimulation of the pyramidal tract, the ventrobasal thalamus and the corpus callosum. Neurons excited by acetylcholine could be segregated into two groups: one encompassing layer Vb and the upper part of layer VI, the other more deeply located at the limit between the cerebral cortex and the subjacent white matter. Neuronal responses to glutamate and nicotine, unlike those to actylcholine were evenly distributed in the cortex. Pyramidal tract neurons had corticothalamic neurons were frequently excited by acetylcholine and were shown to be located with the first group of acetylcholine sensitive neurons. Commissural neurons were rarely excited by acetylcholine and were not restricted to either group. The analysis of neuronal responses to acetylcholine and various agonists (carbachol, nicotine, acetyl-beta-methylcholine, carbamyl-beta-methylcholine, butyrylcholine) and antagonists (atropine, mecamylamine) revealed a prominent but not exclusive muscarine character. It is included (i) that cortical neurons of first somatosensory cortex which are excited by acetylcholine belong to two populations, one consisting, at least in part, of projection neurons (upper group) and the other of interneurons (lower group); (ii) that cortical acetylcholine receptors are of a 'mixed' type strongly weighted toward the muscarinic side.

Acetylcholine↗
PubMed1986

Architecture of apical dendrites in the murine neocortex: dual apical dendritic systems.

A monoclonal antibody (5F9) against microtubule-associated protein 2 is a selective and sensitive marker for neocortical dendrites in the mouse. The marker stains all dendrites. It affords a particularly comprehensive picture of the patterns of arrangements of apical dendrites which are most intensely stained with this antibody. Dual systems of apical dendrites arise from the polymorphic neurons of layer VI, on the one hand, and the pyramidal neurons of layers II-V, on the other. Terminal arborization of the former is concentrated principally at the interface of layers V and IV, while that of the latter is in the molecular layer. Apical dendrites of both systems are grouped into fascicles. In supragranular layers and in upper layer VI-lower layer V, where apical dendrites are most abundant, the fascicles coalesce into septa. These generate a honeycomb-like pattern, subdividing these cortical levels into columnar spaces of approximately 20-40 micron diameter. At the level of layer IV, where the number of apical dendrites is greatly reduced, the fascicles are isolated bundles. These bundles have the form of circular, elliptical or rectangular columns in the primary somatosensory, temporal and frontal regions, respectively. Those in the barrel field are preferentially concentrated in the sides of barrels and the interbarrel septa. The configurations of the dendritic fascicles, particularly the midcortical bundles, may conform to the spatial configuration of investing axons of interneurons.

Animals↗
PubMed1988

Noradrenaline- and vasoactive intestinal peptide-containing neuronal systems in neocortex: functional convergence with contrasting morphology.

Neurotransmitter-specific anatomical techniques have provided a tool to define the morphological constraints within which a given neurotransmitter will exert its cellular actions. Biochemical and electrophysiological approaches have revealed the nature of these cellular actions for several neurotransmitters. Furthermore, by using purified preparations and tissue cultures a certain degree of resolution has been achieved by which the cell type, where a neurotransmitter's effect takes place, can be determined. In this article we review these aspects for noradrenaline and vasoactive intestinal peptide, two neurotransmitters of the cerebral cortex contained within neuronal systems that present strikingly different morphologies. Nevertheless, noradrenaline and vasoactive intestinal peptide share certain cellular actions and can interact synergistically. The experimental evidence accumulated to date indicates that noradrenaline- and vasoactive intestinal peptide-containing neurons can influence three general cell types of the cerebral cortex, i.e. (i) other neurons, (ii) astrocytes and (iii) cells of the vasculature. This diversity in cellular partners supports the notion that noradrenaline and vasoactive intestinal peptide can be released from neurons at conventional synapses as well as at extrasynaptic sites, thus suggesting the co-existence of two modes of release within the same neuron.

Cerebral Cortex↗
PubMed1990

Noradrenaline axon terminals in adult rat neocortex: an immunocytochemical analysis in serial thin sections.

Peroxidase-antiperoxidase electron microscope immunocytochemistry with an antiserum against noradrenaline-glutaraldehyde-protein conjugate was used to identify cortical noradrenaline terminals (axonal varicosities) from the upper layers of the frontal, parietal and occipital cortex in adult rat. A large number of immunostained varicosities were examined in serial thin sections, and compared with a control population of randomly chosen unlabeled terminals from the same sections. Both groups of varicosities were measured and scrutinized for the presence of a junctional complex indicative of synaptic specialization. Cellular elements juxtaposed to the membrane of both types of varicosities were also identified and counted. Noradrenaline varicosities in all three cortical regions averaged 0.65 microns in diameter. In contrast to their unlabeled counterparts, these profiles rarely showed a membrane differentiation characteristic of a synaptic contact (junctional complex). The rare junctional complexes formed by cortical noradrenaline varicosities were invariably symmetrical and almost always found on dendritic shafts. The microenvironment of noradrenaline varicosities also differed, exhibiting a greater number of apposed axonal varicosities and a smaller number of dendritic spines than that of the random population. The proportion of noradrenaline varicosities making a synaptic contact (synaptic incidence) was determined by plotting the incidence of visible junctions as a function of the number of thin sections available for examination. As extrapolated for whole varicosities after linear transformation (double reciprocal plot), this proportion was 17% or 26% depending on the stringency of the criteria used in identifying the junctional complex. The same analysis provided a figure of 98% for the control population. The present study largely confirmed our initial radioautographic characterization of the cortical noradrenaline innervation as a mostly non-junctional system, and also indicated that these varicosities are set in a particular microenvironment. These new data further support the eventuality of a diffuse release of cortical noradrenaline in the extracellular space, compatible with both its neuromodulatory role and multiplicity of actions on diverse cellular targets in the cerebral cortex. The functions assigned to the coeruleocortical noradrenaline system must therefore be viewed as the product of a widespread and ubiquitously distributed neuronal organization characterized by loose intercellular relationships. This system might be capable of selectivity and specificity of action, however, owing to the distribution of its receptors, and in view of intrinsically or extrinsically driven control mechanisms triggered by the release of its own or other transmitters and which may also involve target-initiated feedback mechanisms.

Animals↗
PubMed1990

Augmentation by glycine and blockade by 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX) of responses to excitatory amino acids in slices of rat neocortex.

Responses of neocortical pyramidal cells to excitatory amino acids were recorded intracellularly. Agonists and antagonists were applied electrophoretically from a separate multibarrel pipette and care taken to ensure that the pipette was positioned to evoke optimal responses to N-methyl-D-aspartate (NMDA), or homocysteic acid, before control responses were recorded. Responses to NMDA, but not those to alpha-amino-3-hydroxy-5-methyl-4-isoxazdepropionic acid (AMPA) or quisqualate, were enhanced when glycine was co-applied. Responses to AMPA, quisqualate and NMDA were reduced by 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX) applied either electrophoretically, or in the bathing medium, with responses to quisqualate being the least and those to AMPA being the most sensitive to CNQX. The blockade of NMDA responses by CNQX was selectively reversed by additional glycine confirming that CNQX blocks NMDA receptor-channel complexes at the glycine, rather than at the NMDA site. Under control conditions, responses to glutamate resembled responses to quisqualate, and were relatively insensitive to CNQX, 3-((+/-)-2-carboxypiperazin-4-yl)-propyl-l-phosphonic acid and 2-amino-5-phosphonovalerate, while responses to homocysteic acid resembled responses to NMDA and were blocked by these antagonists. This suggested that homocysteic acid acted at NMDA receptors, while glutamate acted primarily at non-NMDA receptors. However, responses to both glutamate and homocysteic acid were augmented by additional glycine when these transmitter candidates were applied close to a "hot spot" for NMDA receptor activation. The glycine enhancement of responses to glutamate was sensitive to NMDA antagonists, indicating that glutamate can activate NMDA receptors in an intact preparation if glycine levels are high enough.

6-Cyano-7-nitroquinoxaline-2,3-dione↗
PubMed1990

Paroxysmal afterpotentials and role of calcium-dependent potassium conductivity in neuronal activity of strychninized neocortex.

Reactions of cortical suprasylvian gyrus neurons were investigated intracellularly after supracortical strychnine application in immobilized and anaesthetized cats. It was shown that paroxysmal depolarizing shifts of membrane potential could be accompanied by de- and hyperpolarizing afterpotentials. When passing from epileptiform to normal physiological activity, short afterhyperpolarizations, 300-500 ms in duration, were converted into inhibitory postsynaptic potentials which were also accompanied by a decrease in membrane potential. When the frequency of paroxysmal discharge was less than 1 s, prolonged (1-2 s) afterhyperpolarizations were observed; at a higher frequency their summation led to tonic hyperpolarization of the membrane. The ictal discharges were accompanied by postictal hyperpolarizations of up to 30 s duration. The intracellular injection of EGTA blocking Ca2(+)-dependent potassium conductivity eliminated prolonged after- and postictal hyperpolarizations and produced depolarizing afterpotentials and a gradual depolarization of cell membranes. Our results indicate that the development of short hyperpolarizing afterpotentials could be determined by the inhibitory synaptic effects. The activation of Ca2(+)-dependent potassium conductivity caused by the development of prolonged afterhyperpolarizations and postictal polarizations, as well as maintained tonic hyperpolarization of cell membranes. Obviously, the depolarizing afterpotentials are of a non-synaptic origin and can be induced by inward calcium current.

Animals↗
PubMed1991

Prenatal development of the intrinsic neurons of the rat neocortex: a comparative study of the distribution of GABA-immunoreactive cells and the GABAA receptor.

The ontogenesis of cells showing GABA-like immunoreactivity, and the distribution of the immunoreactivity for the GABAA receptor were studied immunocytochemically in the prenatal rat brain. By embryonic day 14, a few GABA-like immunoreactive (GABA-positive) cells scattered at the subpial limit of the marginal zone (primordial plexiform layer) in the lateral part of the developing cortex. GABA-positive cells appeared progressively within the dorsal and medial sectors of the primordial plexiform layer, occupying deeper positions within the layer. The immunoreactivity for the GABAA receptor covered the whole thickness of the primordial plexiform layer. By embryonic day 16, most GABA-positive cells populated three distinct laminar compartments of the developing cortex: the prospective lamina I, the subplate, and the lower part of the intermediate zone. The GABA-positive cells of the lower intermediate zone appeared to be typical of the developing cerebral cortex of the rat: their neuronal nature was assessed immunocytochemically, using monoclonal antibodies against microtubule-associated protein 2, mainly expressed in neuronal somata and dendrites, and against intermediate filament protein vimentin, expressed in glia. The lower intermediate zone contained cells immunoreactive for microtubule-associated protein 2, although the immunostaining was less intense than in the prospective lamina I and the subplate. Preliminary results showed no vimentin-positive cells in the lower intermediate zone. At embryonic day 16, immunoreactivity for the GABAA receptor was present within the prospective lamina 1 and the subplate. Preliminary results showed no vimentin-positive cell in the lower intermediate zone. At embryonic day 16, immunoreactivity for the GABAA receptor was present within the prospective lamina 1 and the subplate, but not in the lower intermediate zone. From embryonic day 18 onwards, the immunostaining for the GABAA receptor labelled, unambiguously, the subplate as a lamina clearly separated from the suprajacent cortical plate. At embryonic day 18, the GABAA receptor started to be expressed within the lower, differentiating part of the cortical plate. Within the cortical plate, the expression of GABA in neural cell perikarya, and the immunostaining for the GABAA receptor, followed a similar spatio-temporal ("inside-out") gradient during pre- and early postnatal stages. Most GABA-positive cells of the lower intermediate zone started to disappear (or stopped the expression of GABA) by embryonic day 20, but some remained until adulthood. A similar time-course was observed for the microtubule-associated protein 2-immunoreactive cell population located at the same level.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1992

Background firing activity in guinea-pig neocortex in vitro.

The background firing activity was recorded extracellularly in experiments on guinea-pig neocortical slices maintained in vitro. The following types of background firing activity were revealed: (i) high regular single spikes (48%), (ii) irregular single spikes (15%), (iii) bursts (7%), (iv) groups (7%), (v) mixed activity where single spikes alternated with bursts or groups (28%). The specific interspike interval distribution and the specific shape of autocorrelogram corresponded to each of these background firing activity types. Furie analysis of autocorrelograms showed periodic components in spike sequences with the maxima at 3, 12, and 28 Hz. When blocking synaptic transmission with 100 mM adenosine, about 70% of the background active cells "fell silent" and the remaining 30% of neurons continued to generate action potentials. The latter seem to be actual spontaneously active neurons, i.e. they were capable of autonomous spike generation. We failed to find any correlation between the type of neuronal firing and the ability of neurons to be spontaneously active. The selective blockade of inhibitory synapses with 100 mM picrotoxine did not practically change the character of background firing activity though the responses to stimulation became epileptic. An important conclusion to emerge from this study is that the background firing activity in cortical slices can include the actual spontaneous discharges related to intrinsic cell properties as well as those concerned with synaptic actions. Furthermore, a small number of spontaneously active neurons seem to be able to synaptically activate twice the number of cells. The inhibitory interneurons did not significantly influence the propagation of excitation with the absence of stimulation.

Adenosine↗
PubMed1992

A comparison of the electrophysiological properties of morphologically identified cells in layers 5B and 6 of the rat neocortex.

In vitro studies performed in mammalian brain slices have shown that cortical neurons differ in their intrinsic membrane properties. In the rodent cortex these properties are related to a specific cell morphology and synaptic connectivity in some cells but not in others. Due to their small size, little is known about the intrinsic membrane properties of layer 6 cells, however, and it is not clear whether cell morphology is related to electrophysiological properties in this layer. We used a combination of intracellular recording and dye-filling to study the electrophysiological and morphological characteristics of layer 6 cells of the rat sensorimotor cortex in vitro and compared their properties to those of large layer 5B pyramidal cells. Our sample of 24 filled and anatomically reconstructed cells in layer 6 confirms previous Golgi studies that showed them to be a morphologically diverse group consisting of regularly and irregularly oriented pyramidal cells and spiny nonpyramidal cells. Regular layer 6 pyramidal cells differed with respect to the length of their apical dendrites and extent of their axonal arborizations, while irregularly oriented pyramidal cells consisted of sideways or inverted pyramidal cells of variable size and morphology. Spiny nonpyramidal cells included bi-tufted and multi-polar cell types that differed in size and extent of dendritic trees. Many layer 6 cells showed long horizontal axon collaterals in layer 6, and an oblique or vertical projection to layer 4. Stimulation with intracellular constant current pulses revealed that the morphological diversity was mirrored by a similar electrophysiological diversity. Most layer 6 cells were capable of firing trains of action potentials characterized by an initial doublet or triplet followed by a train of single spikes (phasic-tonic mode). The majority of layer 6 cells could fire in either a tonic (single spikes only) mode with low strength current input and a phasic-tonic pattern with higher current strengths. A minority fired either always phasic-tonic or tonic-only spike trains. The size and sequence of spike afterpotentials during low-rate repetitive firing was highly variable in layer 6 cells suggesting that the relative importance of ionic currents responsible for spike repolarization and afterpotentials varied from cell to cell. Subthreshold responses showed prominent inward rectification, while hyperpolarizing "sag" was present in most cells tested. In comparison, large layer 5B pyramidal cells fired either phasic-tonic only or both phasic-tonic and tonic patterns. A minority of cells were capable of firing repetitive bursts, while the remainder fired repetitive single spikes.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1993

Fluctuations in pyramid-pyramid excitatory postsynaptic potentials modified by presynaptic firing pattern and postsynaptic membrane potential using paired intracellular recordings in rat neocortex.

Single axon excitatory connections between pairs of neocortical pyramidal neurons were studied using paired intracellular recordings in layers II/III and IV of coronal slices of adult rat somatosensory/motor cortex. Excitatory postsynaptic potentials evoked with different presynaptic firing patterns and at different postsynaptic membrane potentials were compared. Two methods of statistical analysis were used in attempts to determine whether changes in mean excitatory postsynaptic potential amplitude were due to presynaptic or postsynaptic modifications. Analysis of the decrease in mean excitatory postsynaptic potential amplitude associated with increases in presynaptic firing rate were consistent with a change in probability of transmitter release. Paired pulse depression appeared to exhibit both presynaptic and postsynaptic components when the interspike interval was < 10 ms, but could be explained simply by a decrease in probability of release with interspike intervals between 10 and 80 ms. Previous studies had demonstrated that these excitatory postsynaptic potentials are partially mediated by N-methyl-D-aspartate receptors. In contrast to the apparently presynaptic effects of firing pattern, postsynaptic membrane depolarization appeared to produce an increase in quantal amplitude. In addition to this increase at low frequencies, a form of frequency-dependent, self-potentiation involving the recruitment of an additional, longer-latency postsynaptic component occurred at higher presynaptic firing rates. The possibility is discussed that two different mechanisms are involved in the replacement of vesicles at release sites. Over a few tens of milliseconds (paired-pulse depression) availability of releasable transmitter may be determined by the rate of replacement of discharged vesicles from a readily releasable pool of vesicles. Over longer periods of firing at 0.33-2 Hz, the readily releasable pool may become exhausted and require replenishment. Postsynaptic depolarization increases the duration of these excitatory postsynaptic potentials, facilitating summation and enables two components of excitatory postsynaptic potential enhancement at N-methyl-D-aspartate receptor-mediated synapses; one that is present at all firing rates and relates simply to voltage dependent events and one that occurs at higher firing rates and involves a gradual, time dependent event. These data also indicate that the optimal pyramidal firing pattern if another pyramid is to be activated is a tonic, or brief burst pattern at relatively low repetition rates. Long bursts of many presynaptic spikes recruit little that is not activated by pairs of spikes. This situation is in stark contrast to the results obtained in the following paper in which excitatory inputs from pyramids to non-pyramids are described.

Animals↗