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[Inhibitory interneuronal interactions in microregions of the visual cortex].

The interactions between the neighbouring neurones of the visual cortex were studied in the background multineuronal activity of awake cats and of cats anaesthetized with Nembutal. The experiment was carried out on 64 trineuronal microsystems with the method of cross-correlation analysis. Those histograms of cross-correlation were analyzed, in which the parameters and configuration seemed to be determined by inhibitory interneuronal interactions. It is assumed that this inhibition is achieved via recurrent axonal collaterals. The duration and the intensity of inhibitory effect, observed in anaesthetized cats, greater than those in awake animals, may be due to a decrease in the general excitatory synaptic drive and to blocking of the unspecific brain system by Nembutal.

Anesthesia, General↗

Doses of 6-hydroxydopamine sufficient to deplete norepinephrine are not sufficient to decrease plasticity in the visual cortex.

These experiments were designed to test Kasamatsu and Pettigrew's (1979, 1983, and see below) hypothesis that plasticity in the visual cortex requires cortical norepinephrine (NE). Kittens were treated with various doses of intraventricular 6-hydroxydopamine (6-OHDA) or vehicle solution. Cortical NE content was measured with high-performance liquid chromatography with electrochemical detection. We sutured the right eyes of some kittens approximately 6 weeks of age for 1 week and recorded from the left visual cortex of these kittens at the end of the week of suture. We measured the ability of the deprived eye to drive cortical cells in animals that received either 0.2 or 4.8 mg of 6-OHDA, and also in control animals that received only vehicle solution. We concluded that a particular dose of 6-OHDA decreased plasticity if it increased (relative to controls) the ability of the deprived eye to drive cortical cells. Doses of 6-OHDA as small as 0.2 mg were sufficient to produce approximately maximal depletion of NE but did not decrease cortical plasticity. Doses of 4.8 mg or more did decrease cortical plasticity, although not as much as was reported by Kasamatsu and Pettigrew. We conclude that 6-OHDA can alter cortical plasticity but the decrease in plasticity does not result from NE depletion.

Animals↗

Molecular analysis of trkC in the cat visual cortex.

trkC belongs to the trk family of neurotrophin receptors. Several isoforms of trkC have been cloned to date; a full-length catalytic form containing a tyrosine kinase (TK) domain, three full-length isoforms with amino-acid insertions (14, 25, and 39 amino acids) in the TK domain, and five noncatalytic truncated forms that completely lack the TK domain. These isoforms have been studied in several mammalian species, including the pig, rat, mouse, monkey, and human. In this article we report the cloning and sequencing of five trkC isoforms isolated from 30-d postnatal cat visual cortex. The first isoform corresponded to the previously reported full-length trkC transcript containing the 14 amino-acid insert. To search for the presence of other inserts, reverse transcription polymerase chain reaction (RT-PCR) was performed on 30-d postnatal cat visual cortex mRNA using primers that flank the insertion site in the TK domain. Both the isoform containing the 14 amino-acid insert and the isoform lacking any insertion were present in abundant amounts, whereas the other two insert containing isoforms (TK25 and TK39) were much less abundant. The fifth isoform discovered corresponds to the previously reported truncated transcript. Overall, there is a high degree of identity (89-98%) and homology (97-99%) between the cat trkC nucleotide and amino-acid sequences among all mammals. The extracellular juxtamembrane domain was found to be highly divergent among all mammals that have been studied to date. This divergent region also included a proline deletion in the cat trkC sequence. This is the first report of the cloning, sequencing, and RT-PCR analysis of trkC in cat visual cortex, a system extensively studied using anatomical and physiological approaches.

Amino Acid Sequence↗

Anatomical organization of macaque monkey striate visual cortex.

I hope that this review of the internal anatomy of the monkey primary visual cortex makes clear the high degree of specialization that exists in each of the cortical laminae and their constituent neurons. Each lamina is driven by different patterns of relays from the LGN and by different patterns of intrinsic interlaminar projections. The elaborate laminar and intralaminar segregation of efferent neuron arrays suggests that the extraordinary precision of inter- and intralaminar connectivity provides a unique functional role for each set of efferent neurons. The organization of aspinous (presumed inhibitory) local circuit neurons suggests that they are highly specialized, and within each lamina and via interlaminar relays each variety may only accomplish a single, particular task. The cortex neuropil does not give the immediate impression of "random" networks, and if such exist, they must surely be between very tightly determined subgroups of neurons. Clearly a very detailed physiological exploration of V1 is still needed, with new consideration of thalamic axon function, of efferent neuron characteristics, of laminar differences, and of spatial organization of properties within laminae, in order to match known anatomical detail with function. The concept of columnar organization in cortical organization of V1 may eventually be redefined in more complex terms that accurately describe the anatomical and functional parcellation evident in cortical depth and perhaps may link it to a means by which a correlation of different aspects of the visual image is achieved.

Animals↗

Cholinergic modulation of synaptic transmission in the rat visual cortex in vitro.

Cholinergic synaptic modulation in the rat visual cortex was studied using intracellular recordings from slice preparations. A cholinergic agonist, carbachol (CCh), reduced fast excitatory as well as fast and slow inhibitory postsynaptic potentials evoked by white matter stimulation. This effect was antagonized by atropine. CCh perfusion did not reduce glutamate- or gamma-aminobutyric acid-induced depolarizations, suggesting the presynaptic mechanism of the suppression. CCh augmented firing over a long period after transsynaptic stimulation combined with a long depolarizing current pulse, not only due to a decrease in firing accommodation but also due to disinhibition. CCh also induced a large sustained depolarization and bursting of action potentials triggered by tetanic stimulation. These results suggest that cholinergic modulation results in a prolonged increase in neuronal excitability during the late phase of synaptic transmissions at least partly by the mechanism of decreasing inhibitory transmissions, particularly when the synaptic inputs are strongly activated.

Animals↗

A slice preparation preserving the callosal projection to contralateral visual cortex.

Due to the curved path they follow, the visual callosal projections to areas OC1 and OC2 of the rat visual cortex have been inaccessible to studies using brain slices. In this paper we describe a new slice preparation in which a curved cutting blade was used to obtain slices in which callosal fibers projecting to OC1 or OC2 are preserved. Stimulation of the contralateral white matter resulted in EPSPs recorded in layer II/III and V cells of OC2 studied with intracellular recording. Current source density analysis of extracellular field potentials collected in OC1 and OC2 revealed laminar current sink patterns paralleling the laminar distribution of callosal terminations reported by Miller and Vogt (Dev. Brain Res., 14 (1984) 304-309). Exposure of slices to 2 mM kynurenic acid reversibly abolished current sinks in OC1 recorded in response to callosal stimulation indicating that glutamate receptors mediate the response of OC1 to callosal afferent activity. This new slicing technique can be readily adapted to study other systems in the nervous system in which neural processes follow curved trajectories.

Animals↗

Glutamate decarboxylase-immunoreactive terminals of Golgi-impregnated axoaxonic cells and of presumed basket cells in synaptic contact with pyramidal neurons of the cat's visual cortex.

Glutamate decarboxylase (GAD)-immunoreactive varicosities were found around cell bodies of nonimmunoreactive and immunoreactive neurons in the cat's visual cortex; they also occurred along apical dendrites and axon initial segments of pyramidal neurons. By examination in the electron microscope of structures first identified in the light microscope, it was established that the GAD-immunoreactive varicosities were boutons in symmetrical synaptic contact with pyramidal cells in layers II-IV. More than 90% of 142 boutons surrounding the cell bodies of 20 pyramidal neurons were immunoreactive for GAD. Since such a high proportion of the axosomatic boutons are GAD-immunoreactive, it is likely that the terminals of basket cells are included in this population and so the basket cell probably uses gamma-aminobutyrate as a transmitter, as suggested by previous authors. Almost all the 68 boutons in symmetrical contact with the axon initial segments of six pyramidal neurons could be shown to be GAD-immunoreactive, which makes it very likely that the boutons of axoaxonic cells contain GAD-immunoreactivity. This was established unequivocally for an individual Golgi-impregnated axoaxonic cell by combining Golgi impregnation and immunocytochemistry in the same sections: A Golgi-impregnated axoaxonic cell whose cell body was in layer II gave rise to numerous terminal segments, some of which were examined in the electron microscope after gold-toning. These boutons were in synaptic contact with axon initial segments and not only contained the Golgi precipitate but were also immunoreactive for GAD. It is concluded that the axoaxonic cell in the visual cortex uses gamma-aminobutyrate as a transmitter. An individual axoaxonic cell in layer II/III was filled with horseradish peroxidase by intracellular iontophoresis. The very extensive local axonal field was composed of 330 terminal bouton rows in layer II/III and a sparse descending collateral projection to infragranular layers. A computer-assisted reconstruction of the axonal field in three dimensions revealed the following: The main output of the cell is to pyramidal neurons that lie deeper than the soma; the axonal arborization occupies an area of 400 micron in the anteroposterior axis and extends 200 micron along the mediolateral axis; the terminal bouton rows in layer II/III form clusters about 50 micron wide running approximately at right angles to the border between areas 17 and 18, with an intercluster interval of about 100 micron. These findings suggest that the terminals of an individual axoaxonic cell could be contained within one ocular dominance column but that there may be inhomogeneities in the weighting of the axoaxonic input to pyramidal cells in the supragranular layers.

Animals↗

[Temporal signal summcation by at visual cortex neurons].

The summation temporal of signals in neurons of field 17 of the visual cortex was studied in immobilized and unanaesthetized cats. The unit responses to optimal local light stimuli were recorded under conditions of light adaptation. Critical duration of the summation temporal in different units was 5-100 ms, with mean value equal to 31.45 +/- 5.67 ms. Neurons in the central part of the field differed from the peripheral ones by a shorter critical summation temporal, a lower background discharge frequency and a smaller duration of the first burst in the response. In neurons with simple receptive fields the summation temporal was considerably shorter than in complex neurons. The obtained results are compared with data from literature on neurons of cat retina and lateral geniculate nucleus and discussed with respect to division of ascending afferent visual projections into X and Y (1a and 1b) groups.

Adaptation, Physiological↗

[Morphologic characteristics of neuroglioform cells in the visual cortex of various mammals (rat, guinea pig, alticola and cat). A Golgi study].

Neuroglioform cells have been studied and compared in visual cortex of rat, guinea pig, alticola and cat in Golgi-material. These neurons possess a distinct morphological pattern, a dense and strictly local dendritic field with many short, fine and radiated dendrites which highly branch near the soma and a very dense axonal plexus of thin, varicose and sinuous branches. Neuroglioform cells were identified in all species and in all laminae of visual cortex. Some features, the size of the soma and the size of the axonal arborization vary among the species. We found small, medium sized and large somata. In general, the axons are confined to the vicinity of the dendritic field, but they can extend to considerable distances beyond it. The form of the soma and the dendritic surface were found to be typical for special species, so the form of the soma in guinea pig neuroglioform cells can be of different size, especially unregularly. In rodents the dendrites bear spines, but the dendrites of the cat are smooth.

Animals↗

Developmental changes in the susceptibility to long-term potentiation of neurones in rat visual cortex slices.

We investigated with intracellular recordings from rat visual cortex slices whether the susceptibility to undergo long-term potentiation (LTP) is age-dependent and whether it is correlated with the expression of synaptic responses mediated by N-methyl-D-aspartate (NMDA) receptors. Test and tetanic stimuli were applied to the white matter and post-tetanic modifications of the amplitude of postsynaptic potentials (PSPs) were assessed in regular spiking cells of supragranular layers. At 2 weeks of age, the amplitudes of early (8-10 ms post-stimulus) and late (20 ms post-stimulus) PSP-components increased after tetanic stimulation to 137.1 +/- 13.4% and 141.3 +/- 12.1% of the pretetanic controls, respectively. At 3 weeks, potentiation of both PSP-components was less pronounced but still significant, the late component being on average more potentiated than the early one. At 4 weeks, PSPs were no longer potentiated. Bath application of 25 microM DL-2-amino-5-phosphonovalerate (APV), an NMDA receptor antagonist, blocked LTP induction both at 2 and at 3 weeks. We also studied developmental changes of two synaptic responses known to influence the susceptibility of cortical neurones to LTP, the NMDA receptor-mediated excitatory PSP (EPSP) and the initial inhibitory PSP (iIPSP). The amplitude of the APV-sensitive EPSP decreased with age and reached adult values in 4-week-old animals. The iIPSPs were pronounced already at 2 weeks and showed no marked change during further development. The results suggest a close correlation between the susceptibility to undergo LTP and the extent to which NMDA receptor-gated conductances contribute to the synaptic response.(ABSTRACT TRUNCATED AT 250 WORDS)

2-Amino-5-phosphonovalerate↗

Changing patterns of synaptic input to subplate and cortical plate during development of visual cortex.

1. The development of excitatory activation in the visual cortex was studied in fetal and neonatal cats. During fetal and neonatal life, the immature cerebral cortex (the cortical plate) is sandwiched between two synaptic zones: the marginal zone above, and an area just below the cortical plate, the subplate. The subplate is transient and disappears by approximately 2 mo postnatal. Here we have investigated whether the subplate and the cortical plate receive functional synaptic inputs in the fetus, and when the adultlike pattern of excitatory synaptic input to the cortical plate appears during development. 2. Extracellular field potential recording to electrical stimulation of the optic radiation was performed in slices of cerebral cortex maintained in vitro. Laminar profiles of field potentials were converted by the current-source density (CSD) method to identify the spatial and temporal distribution of neuronal excitation within the subplate and the cortical plate. 3. Between embryonic day 47 (E47) and postnatal day 28 (P28; birth, E65), age-related changes occur in the pattern of synaptic activation of neurons in the cortical plate and the subplate. Early in development, at E47, E57, and P0, short-latency (probably monosynaptic) excitation is most obvious in the subplate, and longer latency (presumably polysynaptic) excitation can be seen in the cortical plate. Synaptic excitation in the subplate is no longer apparent at P21 and P28, a time when cell migration is finally complete and the cortical layers have formed. By contrast, excitation in the cortical plate is prominent in postnatal animals, and the temporal and spatial pattern has changed. 4. The adultlike sequence of synaptic activation in the different cortical layers can be seen by P28. It differs from earlier ages in several respects. First, short-latency (probably monosynaptic) excitation can be detected in cortical layer 4. Second, multisynaptic, long-lasting activation is present in layers 2/3 and 5. 5. Our results show that the subplate zone, known from anatomic studies to be a synaptic neurophil during development, receives functional excitatory inputs from axons that course in the developing white matter. Because the only mature neurons present in this zone are the subplate neurons, we conclude that subplate neurons are the principal, if not the exclusive, recipients of this input. The results suggest further that the excitation in the subplate in turn is relayed to neurons of the cortical plate via axon collaterals of subplate neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Aging↗

[Focal evoked potentials in the rabbit visual cortex: density analysis of current sources].

Focal evoked potentials were elicited in the rabbit visual cortex by punctiform light stimuli and analyzed by the current source density technique. They contained two main components. The first component was generated by local sink at depths form 0.6 to 1.0 mm (layer IV) with 30 ms latency and peak time about 50 ms. The second one was generated by less local sink at depths form 0.2-0.3 to 1.3-1.5 mm (layers III-VI) with peak time 90-100 ms. These two sinks are considered as active and indicating the localization of depolarizing synapses. Passive sources are dissipated around the zone of the active sinks.

Animals↗

Fast and slow components of unitary EPSCs on stellate cells elicited by focal stimulation in slices of rat visual cortex.

1. Voltage and current recordings were made from visually identified non-pyramidal neurones in slices of layer IV of rat primary visual cortex using the whole-cell configuration of the patch clamp technique. These neurones are characterized by a high input resistance (0.5-2 G omega) and a non-adaptive behaviour of action potential frequency following depolarizing current injection, which suggests that they are stellate cells. 2. Excitatory postsynaptic currents (EPSCs) were recorded from these neurones during focal stimulation of neighbouring cells by a second patch pipette, the tip of which was placed on the soma of the stimulated cell. The response amplitude as a function of stimulus strength showed a sharp increase at a critical stimulus strength suggesting that stimulus-evoked currents represent unitary EPSCs. 3. In most cases the latencies of stimulus-evoked EPSCs were unimodally distributed with means in the range of 2.1-3.6 ms. In some experiments two peaks were seen in the distribution of latencies. The EPSC rise times, measured as the time from 20 to 80% peak amplitude, fell into a distribution ranging from 0.1 to 0.8 ms with a peak at 0.2 ms. The EPSC decay time course at -70 mV membrane potential was fitted by a single exponential with a time constant of 2.39 +/- 0.99 ms (mean +/- S.D.). The rise and decay times were independent of EPSC peak amplitudes. 4. The peak amplitude of successive unitary EPSCs, elicited by a constant stimulus, fluctuated at random. At a holding potential of -70 mV the peak amplitudes varied between 5 and 90 pA. In two out of ten cells the histogram of peak amplitudes could be well fitted by the sum of several equidistant Gaussians with a peak distance of around 10 pA. This suggests that the quantal conductance change underlying the peak current fluctuations is of the order of 100 pS. 5. At membrane potentials more positive than -70 mV the decay of stimulus-evoked EPSCs showed two components with very different time courses. In standard extracellular solution the current-voltage (I-V) relation for the fast component was almost linear whereas the slow component showed a J-shaped I-V relation with a region of negative slope conductance between -30 and -70 mV.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

The course and termination of corticothalamic fibres arising in the visual cortex of the rat.

Corticothalamic axons have been studied in adult Lister hooded rats with single or dual injections of tracers into the visual cortex. Labelled axons leave medial and lateral injection sites in separate or partially overlapping bundles along parallel trajectories in the subcortical white matter. In the internal capsule they converge and both bundles enter roughly the same sector of the thalamic reticular nucleus (TRN). Their reticular terminal fields, however, differ. Axons from a medial injection site innervate more lateral parts of the TRN than do the axons from lateral injection sites. The most medial third of the TRN is not innervated from area 17 but receives a topographically arranged input from peristriate cortex (Crabtree and Killackey, 1989, Eur. J. Neurosci., 1, 94-109; Coleman and Mitrofanis, 1996, Eur. J. Neurosci., 8, 388-404). The two groups of axons then separate in the dorsal thalamus, axons from medial parts of visual cortex turning caudally into lateral regions of the lateral geniculate nucleus, whereas fibres from more lateral cortex continue into medial parts of the nucleus. Connolly and van Essen (1984, J. Comp. Neurol., 226, 544-564) and Nelson and LeVay (1985, J. Comp. Neurol., 240, 322-330) have shown that in the geniculocortical pathway the two groups of fibres cross over in the subcortical white matter, probably in the region of the subplate. We show that the corticothalamic pathway also has a crossing, but it occurs in, or close to, the diencephalon itself, in the region of the perireticular nucleus. This result suggests that each of these pathways, the geniculocortical and the corticogeniculate, may undergo reorganization within distinct cerebral zones, one diencephalic for the corticothalamic axons and the other telencephalic for the thalamocortical axons.

Animals↗

Synaptogenesis in the primary visual cortex of the tree shrew (Tupaia belangeri).

The primary visual cortex of the tree shrew is characterized by the lack of ocular dominance columns. The two eyes are represented in sublayers of laminae 3 and 4. In an earlier study using the transneuronal transport we observed that the geniculate afferents from the two eyes do not initially overlap and then segregate into their appropriate sublaminae. The final distribution pattern can already be observed during the early postnatal period. Since segregation and elimination of afferent terminal branches do not seem to take place, we wanted to investigate whether or not an overproduction of synapses can be observed as in several other animals. We examined layers 3B, 3C, 4A, and 4B, which receive afferents from the retina via the lateral geniculate nucleus, from P5 to maturity by using the electron microscope. The brain tissue was excised in the region where the central vision is represented in adult animals. Then we determined the density of synapses per 100 microns 2 neuropil for each of the four sublayers at the ages P5, P15, P19, P23, P31, and P42 and in the adult animal (AD). In determining the neuropil we measured the size of two additional compartments, i.e., the compartments consisting of perikarya and of blood vessels. At a higher resolution we determined the fraction of Gray type I and type II synapses in each sublamina and in each developmental stage. The size of the neuropil increases from 57% at P5 to 81% in AD whereas the compartment of perikarya decreases from 42% to 15% and the compartment of blood vessels increases from 1.3% to 3.9%. The synaptic density starts with very low values (3.5/100 microns 2) at P5. Then it increases rapidly and attains a maximal rate of increase during the period of eyelid opening. After this period the increase is slowed down and approaches the adult value (12.5/100 microns 2) slowly. An overproduction of synapses could not be observed. The percentage of type I and type II synapses also changes during this period. The fraction of type I synapses amounts to 73% at P5 and increases to 92% in AD. The increase in density of type I synapses is continuous and does not show any sign of overproduction. The density of type II synapses rapidly reaches it final value and then remains constant. Possibly there is a slight overproduction during the period of eyelid opening.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Dynamics of orientation selectivity in the primary visual cortex and the importance of cortical inhibition.

To test theories of orientation selectivity in primary visual cortex (V1), we have done experiments to measure the dynamics of orientation tuning of single neurons in the V1 cortex of macaque monkeys. Based on our dynamics results, we propose that a V1 cell's orientation selectivity is generated mainly by both tuned enhancement and global suppression. Enhancement near the preferred orientation is probably caused by feed-forward input from LGN (plus amplification by cortical-cortical interaction). Global suppression could be supplied by cortical inhibition. Additionally, in about 1/3 of V1 neurons (usually the most sharply tuned) there is tuned suppression, centered near the cell's preferred orientation but broader than tuned enhancement. These mechanisms also can explain important features of steady-state selectivity in the V1 neuron population. Furthermore, similar neuronal mechanisms may be used generally throughout the cerebral cortex.

Animals↗

Laminar differences in receptive field properties of cells in cat primary visual cortex.

1. Cells in area 17 of the cat visual cortex were studied with a view towards correlating receptive field properties with layering. A number of receptive field parameters were measured for all units, and nearly every unit was marked with a microlesion to determine accurately the layer in which it was found.2. Cells were defined as simple or complex by mapping with stationary stimuli, using the criteria of Hubel & Wiesel (1962). Complex cells fell into two groups: those that showed summation for increased slit length (standard complex) and those that did not (special complex).3. The simple cells were located in the deep part of layer 3, in layer 4, and in layer 6. This corresponds to the distribution of afferents from the dorsal layers of the lateral geniculate nucleus. In these cortical layers the simple cells differed primarily with respect to their receptive field size, cells in layer 4 having the smallest, layer 3 intermediate, and layer 6 the largest fields. Layer 4 was the only layer in which simple cells showed end-inhibition (a reduction in response to slits extending beyond the excitatory portion of the receptive field).4. The standard complex cells were found in all layers, but were quite scarce in layer 4. As with the simple cells, field size varied with layer: in layer 2+3 they had small to intermediate field sizes, in layer 5 intermediate, and in layer 6 very large. Layer 6 cells showed summation for slits of increased length up to very large values, and responded best when the slits were centred in the receptive field. The only standard complex cells that showed end-inhibition were those in layer 2+3, and these were similar to the layer 4 simple cells in terms of proportion of end-inhibited units and degree of end-inhibition.5. The special complex cells, originally described by Palmer & Rosenquist (1974), were found in two tiers: the upper one at the layer 3/layer 4 border and the lower one in layer 5. They were different from the standard complex cells in having a high spontaneous activity, high velocity preference, and large fields which were similar in size (at a given eccentricity) from one cell to the next. Many showed reduced response to slits of increasing length, even for slits that did not extend beyond the borders of the responsive region.6. Cells in layer 6 (the origin of the corticogeniculate projection) were antidromically activated from the lateral geniculate nucleus. The antidromically activated units included both simple and complex cells, and they had the long receptive fields characteristic of the overall population of cells in layer 6.7. The results showed that there are different types of simple and complex cells, and that cells in different layers have different properties. Taken together with their differences in site of projection, this demonstrates that the anatomical lamination pattern is reflected in functional differences between cells in different layers.

Action Potentials↗

[The influence of the posterior hypothalamus on the visual cortex in different states of the reticular formation].

In alert rabbits, stimulation of posterior hypothalamus induced short-latency responses in the visual cortex and affected the formation of the EPs to light. Depending on the intervals between the stimuli, either initial extinction of the EPs (1-15 msec) or subsequent selective facilitation of its positive component with simultaneous depression of negative (20-100 msec) one, or complete recovery of the response (200-300 msec), were observed. Aminazine and benactizine made the effect of the posterior hypothalamic stimulation on visual EPs to light less obvious and changed its dynamics. Other findings suggested an inhibitory effect of the mesencephalic RF on the activity of hypothalamo-cortical input. A role of phasic mechanism of hypothalamic control in realization of visual cortex's perceptive function, is discussed.

Animals↗