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Organization of primary visual cortex (area 17) in the ferret.

Anatomical and electrophysiological mapping techniques were used to determine topographic organization and arrangement of ocular dominance columns in the primary visual cortex of ferrets. From its border with area 18 on the posterior lateral gyrus, area 17 extends around the caudal pole of the hemisphere and over the splenial gyrus to the caudal bank of the splenial sulcus. The visuotopic map is oriented with the isoazimuth lines approximately parallel to the long axis of the posterior lateral gyrus and the isoelevation lines approximately perpendicular to the isoazimuths. Central azimuths are represented on the posterior lateral gyrus and peripheral azimuths are represented on the splenial gyrus; the inferior visual field maps medially and the superior visual field maps laterally. As in other species, the representation of the central visual field is expanded. The ferret has a considerable degree of binocular vision. Receptive fields driven through the ipsilateral eye extended more than 20 degrees into the contralateral visual field. Within the region of area 17 corresponding to the binocular portion of the visual field, tritiated proline injected into one eye transneuronally labelled an ipsilateral projection as a series of patchy bands roughly complementary to gaps in the labelled contralateral projection. Physiological ocular dominance columns were evident as well in that neurons and groups of neurons recorded in this region showed clustered ocular dominance preferences. Most single neurons studied were binocularly driven.

Animals

Development of orientation selectivity in the primary visual cortex of normally and dark reared kittens. I. Kinetics.

Kinetics of the development of orientation tuning are inferred from quantitative analysis of extracellular recordings in the primary visual cortex of normally and dark reared kittens. 712 visual cells were classified in three functional groups: a) non-specific cells, and b) immature cells which are not as orientation selective as c) specific cells. Power regression and covariance analysis indicate that the "critical period" begins before 19 days and that the kinetics of the immature pool are the same in both rearing conditions. A catenary process of development of orientation selectivity is proposed, the immature compartment being a transit pool between non-specific and specific cells. Two sequential stages occur: 1) the realisation of an intrinsic programme of maturation, by which cortical specificity appears at eye opening and increases independently of visual experience 2) a phase of "epigenesis" beginning at 19 days, during which functional modification depends on visual experience.

Animals

[Persistence of orientation-selective cells of the primary visual cortex in kittens enucleated unilaterally at birth and reared in darkness].

In kittens dark reared (6 weeks old) orientation selective cells are no longer recorded in the primary visual cortex, while in kittens of same age, enucleated at birth unilaterally and reared in identical conditions, 30% of visual cortical cells are shown to be orientation selective and in addition respond preferentially to horizontal and vertical orientations.

Animals

Anatomical organization of the primary visual cortex (area 17) of the cat. A comparison with area 17 of the macaque monkey.

Golgi and axonal transport techniques have been used to examine the organization of neurons within primary visual cortex, area 17, of the cat. This organization has been compared to that of the primate cortical area 17 as described in previous studies and it is discussed in relationship to the distribution of afferents from the dorsal lateral geniculate nucleus (dLGN). The visual cortex of the cat and monkey show strong similarities in the laminar positions of neurons projecting extrinsically and also in the restriction of spiny stellate neurons to a central lamina (lamina 4) receiving input from the dLGN. However, lamina 4B in the monkey, which contains spiny stellate neurons but does not receive direct input from the dLGN, has no direct counterpart in cat area 17. Axon projections of spiny stellate neurons in the other divisions of lamina 4 differ in cat and monkey: the small, closely packed neurons in the lowermost division of lamina 4 (4B in the cat, 4Cbeta in the monkey) project chiefly within lamina 4 in the cat whereas in the monkey they have a strong projection to lamina 3. In the cat, spiny stellate neurons of lamina 4A project upon lamina 3 whereas in the monkey those in the apparently equivalent zone, 4Calpha, project upon lamina 4B. Most non-spiny stellate neurons examined have precisely organized interlaminar axonal projections which differ from the axon trajectories of neighboring spiny neurons.

Animals

Histological asymmetry in the primary visual cortex of the rat: implications for mechanisms of cerebral asymmetry.

The present study was designed to specify the contributions of various histological parameters to hemispheric asymmetry of architectonic areas. It was found that the primary visual cortex of the rat is asymmetrical in volume, and that the asymmetry reflects side differences in the number of neurons. The implications of this finding for the understanding of mechanisms involved in the production of brain asymmetries are discussed.

Animals

Evidence for excitatory amino acid neurotransmitters in the geniculo-cortical pathway and local projections within rat primary visual cortex.

To examine the organization of axon collaterals of neurons that selectively take up and transport excitatory amino acids, we have used retrograde tracing with D-[3H]Aspartate after injections into different layers of rat primary visual cortex. The results show cells in the lateral geniculate nucleus retrogradely labeled from the cortex. Additional topographically precise input to the thalamic recipient layer 4 originates from neurons in the visual cortex lying in layers 2/3, 5 and 6. These inputs are reciprocated by point-to-point projections from layer 4. Layer 2/3 cells project to layers 5 and 6 in columnar fashion. Putative excitatory input to layer 2/3 originates from a vertical column of cells in layer 5 and the middle of layer 6. In addition layer 2/3 receives input via horizontal collaterals of topographically distant upper layer neurons, from more widespread projections in lower layer 6, and from very widespread projections of cells at the layer 5/6 border. Cells in the depth of layer 5 also distribute collaterals within layers 5 and 6. Our findings provide anatomical evidence that the geniculo-cortical pathway in the mammalian visual system may use excitatory amino acid transmitters. In addition, the results support the notion that most long range connections that link distant points of the topographic map are excitatory.

Animals

Functional organization of primary visual cortex in the mink (Mustela vison), and a comparison with the cat.

The functional organization of geniculocortical afferents and the visual responses of neurons in primary visual cortex (area 17) were studied in barbiturate-anesthetized, paralyzed minks and cats. Responses of the afferents were studied after silencing intrinsic cortical activity with injections of kainic acid. In both species, afferents were segregated into patches on the basis of eye of origin. In the mink, but not in the cat, there was a further segregation on the basis of center type, with on- and off-center afferents terminating in alternating, partially overlapping patches. The visual responses of cortical neurons in the mink showed many similarities to those in the cat. Nearly all units were orientation-selective, and there was a columnar organization for preferred orientation. Many units were selective for one direction of movement. Within the binocular segment of cortex, although many units could be driven from either eye, there was a marked bias toward the contralateral eye compared to the cat. There was a columnar system for ocular dominance, but contralateral eye columns were wider than ipsilateral. In both species, a quantitative study was made of the responses of cortical neurons to stationary, flashing slits as a function of position in the receptive field. In the mink, and less clearly in the cat, units could be identified as simple or complex on the basis of the spatial separation or overlap of "on" and "off" discharge zones. In both species, simple cells were found most commonly in layers IV and VI, while layer V contained the greatest proportion of complex cells. The relative strengths of the on and off discharges of single cells were also measured. In the mink, many units gave better overall responses to the on or off phase of the stimulus, and 15% showed a strong (greater than 9:1) preference for one or the other, compared to 4% in the cat. In the mink, units with a common preference for the on or off phase of stationary stimuli were arranged in columnar aggregates, a feature of cortical organization that was not found in the cat. These columns probably result from the partial segregation of on-center and off-center geniculate afferents within layers IV and VI of the mink's cortex. On-dominated columns were, however, wider or more numerous than off-dominated columns.

Animals

Image analysis of Nissl-stained neuronal perikarya in the primary visual cortex of the rat: automatic detection and segmentation of neuronal profiles with nuclei and nucleoli.

An image analysing procedure for the morphometric characterization of cortical neurons in Nissl-stained brain sections is described. It consists of the automatic detection of cellular profiles and their compartments: cytoplasm, nucleus and nucleolus. The algorithm was designed to cope with the large morphological spectrum of cortical perikarya (e.g. geometrical properties of perikarya, staining intensities of cell compartments and nucleo-plasmic area-ratio) including pyramidal (Golgi-category I) and non-pyramidal (Golgi-category II) neurons. Clusters of cells were separated and non-neuronal structures (e.g. glia, endothelial cells) as well as tangential, non-nucleolated sections through neuronal perikarya recognized and excluded from further analysis without requiring interactive procedures. The performance of the profile recognition procedure was evaluated using 426 nucleolated and non-nucleolated profiles of different types of neurons in the primary visual cortex of the rat. Nucleolated profiles were recognized as such with a 91% accuracy, non-nucleolated profiles were rejected correctly in 90% of cases. After automatic segmentation and selection of nucleolated neuronal profiles from the microscopic field, a large set of quantitative morphological features including geometrical, densitometrical and textural parameters can be measured using high power light microscopy. This permits quantitative morphometric characterization of different neuronal types. This procedure is the first part of a system for the automatic classification of Nissl-stained cortical neurons.

Algorithms

Texture segregation is processed by primary visual cortex in man and monkey. Evidence from VEP experiments.

We investigated whether the process of texture segregation can be allocated to a specific visual cortical area. We designed a stimulus to reveal the presence of a mechanism, which is specifically sensitive to a checkerboard, that is solely defined by textures segregating due to orientation differences of the constituting line segments. We recorded evoked potentials to this stimulus in man and awake monkey. A difference component, signalling texture segregation sensitivity, could be recorded from both types of subjects. Its presence depended on the spatial extent of the textures, in a manner correlating with the perceptibility of the checkerboard. This difference response could be localized in primary visual cortex by means of equivalent dipole estimations.

Animals

[Double orientation tuning of neurons of the primary visual cortex of the cat at various levels of alertness].

Orientation tuning (OT) of 225 visual cortex neurons was studied in immobilized cats by their responses to flashing light bars. It was found that 43% of neurons had monomodal OT and preferred the horizontal and vertical orientations, while 57% of neurons had double OT, i.e. they had main preferred orientation (PO) and additional PO (aPO). The mean angle between PO and aPO was equal to 71.4 +/- 2.4 degrees. In half of the cases the second maximum of OT was equal to the first one (mean: 0.7 +/- 0.03 from PO). Orientation characteristics of PO and aPO were practically identical. Under light and middle levels of narcosis half of the neurons with double OT became monomodal and 12% of monomodal neurons receive bimodal OT. Monomodal neurons had more often simple RF and invariance of OT to narcosis. The neurons with double OT had simple and complex RF equally often and their OT changed in narcosis. It is supposed that the neurons with double OT can be an angle and line cross detectors. Monomodal neurons may be the stable bench mark system of orientation coordinates. Interaction of these neuronal systems permits conducting an effective analysis of image features in the primary visual cortex.

Anesthesia, General

Where practice makes perfect in texture discrimination: evidence for primary visual cortex plasticity.

In terms of functional anatomy, where does learning occur when, for a basic visual discrimination task, performance improves with practice (perceptual learning)? We report remarkable long-term learning in a simple texture discrimination task where learning is specific for retinal input. This learning is (i) local (in a retinotopic sense), (ii) orientation specific but asymmetric (it is specific for background but not for target-element orientation), and (iii) strongly monocular (there is little interocular transfer of learning). Our results suggest that learning involves experience-dependent changes at a level of the visual system where monocularity and the retinotopic organization of the visual input are still retained and where different orientations are processed separately. These results can be interpreted in terms of local plasticity induced by retinal input in early visual processing in human adults, presumably at the level of orientation-gradient sensitive cells in primary visual cortex.

Discrimination, Psychological

[Primary visual cortex response recovery cycles caused by pharmacologic and electrical effects on the monoaminergic systems of the rat brain].

The recovery cycles (RC) of primary responses in the visual cortex were studied in alert rats following the electrical stimulation of raphe nuclei and locus coeruleus as well as following pharmacological and complex (electrical and pharmacological) actions on serotonin- and noradrenergic brain systems. It was found that both electrical and pharmacological actions on the above monoaminergic systems reduce the duration and depth of relative non-reactivity phase of RC. Complex (electrical and pharmacological) action leads to heterodirectional changes in RC. The character of these changes reflects the reciprocity existing in functional interrelations between serotonin- and noradrenergic brain systems. The probable connection of the found effects with memory mechanisms is discussed.

5-Hydroxytryptophan

The pharmacology of synapses formed by identified corticocollicular neurons in primary cultures of rat visual cortex.

Primary cultures of neurons from the visual cortex of 7-10-d-old Long Evans rats were used to study the pharmacology of synaptic transmission. Dissociated cells were grown either in mass cultures, which contained 8000-10,000 neurons, or in miniature island cultures of 50-100 cells. Prior to dissociation, cells in layer V of visual cortex that project to the superior colliculus were labeled in vivo by retrograde transport of fluorescent latex microspheres-a permanent fluorescent marker. After 2 d to 8 weeks in culture, labeled neurons were identified by epifluorescent illumination, and electrophysiological recordings were obtained from a labeled cell and, simultaneously, from a nearby unlabeled neuron in the same field of view. The 2 neurons were stimulated sequentially by current injection and the pharmacology of evoked postsynaptic potentials (PSPs) was investigated. In mass cultures, relatively few pairs of neurons from which we recorded were synaptically connected, although nearly every cell exhibited abundant spontaneous EPSPs and IPSPs. Neurons grown on island cultures generally did not exhibit spontaneous synaptic activity; however, stimulation of one of the cells in a pair frequently elicited a short-latency PSP in the follower neuron. Retrogradely labeled corticocollicular neurons produced only excitatory PSPs in follower cells, while unlabeled neurons were either excitatory or inhibitory. Three antagonists of excitatory amino acid receptors, kynurenic acid, piperidine dicarboxylic acid, and gamma-D-glutamylglycine, completely blocked EPSPs produced by labeled corticocollicular neurons, as well as EPSPs produced by nearly all of the unlabeled excitatory cells. We have previously shown that these compounds block both N-methyl-D-aspartate (NMDA)-type and non-NMDA receptors on cultured cortical neurons (Huettner and Baughman, 1986). The specific NMDA receptor antagonist 2-amino-5-phosphonovaleric acid (APV) did not alter short-latency EPSPs recorded in 1 mM Mg2+, but did reduce longer-latency EPSPs polysynaptic activity. Since responses mediated by the NMDA receptor are known to be antagonized by Mg2+ (Mayer and Westbrook, 1985), we perfused cultures with Mg2+-free medium and found that the falling phase of some monosynaptic EPSPs was prolonged. Addition of APV to Mg2+-free medium reduced the duration of the falling phase of EPSPs such that they returned to the time course obtained in 1 mM Mg2+.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

Dendritic morphology and axon collaterals of corticotectal, corticopontine, and callosal neurons in layer V of primary visual cortex of the hooded rat.

Recent evidence indicates that corticotectal neurons belong to only one of the three morphological classes of pyramidal cells in layer V. The present study compares the dendritic morphology and axon collaterals of corticotectal, corticopontine, and layer V callosal neurons by using techniques based on the retrograde transport of horseradish peroxidase and fluorescent dyes as well as in vitro intracellular dye injections. Our results indicate that corticotectal and corticopontine neurons are located predominantly in the upper middle part of layer V. These neurons have medium to large somas with 5 or 6 primary basal dendrites and a single apical dendrite ascending to layer I. Approximately 60% of these cells send axon collaterals to both the superior colliculus and the pons. In contrast, callosal neurons form a heterogeneous group. In general, they have small pyramidal or ovoid cell bodies which give rise to 3 or 4 primary basal dendrites. Many cells have an apical dendrite that bifurcates and terminates in layer V or IV. We find that callosal neurons do not send an axon collateral to either the superior colliculus or the pons. We conclude that the corticotectal and corticopontine systems are similar in their intralaminar distribution, dendritic morphology, and pattern of axon collaterals, whereas the callosal system differs in these characteristics.

Animals

The origin of efferent pathways from the primary visual cortex, area 17, of the macaque monkey as shown by retrograde transport of horseradish peroxidase.

The retrograde transport of horseradish peroxidase has been used to identify efferent cells in area 17 of the macaque. Cells projecting to the lateral geniculate nucleus are small to medium sized pyramidal neurons with somata in lamina 6 and the adjacent white matter. The projection to the parvocellular division arises preferentially from the upper half of lamina 6, while that to the magnocellular division arises preferentially from the lower part of the lamina. The projection to both superior colliculus and inferior pulvinar arises from all sizes of pyramidal neurons lying in lamina 58 (Lund and Boothe, '75); at least pyramidal neurons of lamina 5B send collateral axon branches to both destinations. Injections with extensive spread of horseradish peroxidase show that many cells of lamina 4B and the large pyramidal neurons of upper lamina 6 also project extrinsically but their terminal sites have not been identified. Other studies have indicated that cells of laminae 2 and 3 project to areas 18 and 19. Therefore every lamina of the visual cortex, with the exception of those receiving a direct thalamic input, contains cells projecting extrinsically. Further, each lamina projects to a different destination and from Golgi studies can be shown to contain cells with specific patterns of dendritic branching which relate to the distribution of thalamic afferents and to the patterns of intracortical connections. These findings emphasise the significance of the horizontal organisation of the cortex with relation to the flow of information through it and contrast with the current concept of columnar organisation shown in physiological studies.

Animals

Anatomical organization of primary visual cortex (area 17) in the ferret.

The present report describes the intrinsic and extrinsic cortical connectivity of striate cortex (area 17) in the ferret. Injections of horseradish peroxidase demonstrate periodic intrinsic connections over an extent of 2.5-3.0 mm, mainly in the supragranular layers but also occurring secondarily in layer 5. These connections have a stripelike configuration, with a center-to-center spacing of 0.5-0.7 mm. Their laminar distribution and stripelike configuration resemble the pattern in the cat (Gilbert and Wiesel, '83), another member of the carnivore family, but not that in monkeys. In both macaque and squirrel monkeys, these connections have a bilaminar distribution in layers 2-3 and 4B, and a more complicated latticelike geometry (Rockland and Lund, '83). Their interperiod spacing, of about 0.5 mm, however, is relatively constant across species. Extrinsic connections in the ferret link striate cortex with territories probably homologous to feline areas 18 and 19, and to the suprasylvian region. Callosal connections extend on the lateral surface about 1.5 mm into area 17 and 4.0 mm into area 18 beyond their common border. There are homotopical connections between striate cortices and heterotopical connections from at least areas 18 and 19 to contralateral area 17. In addition to gray matter connections, intracortical injections also result in labeled interstitial neurons in the subgriseal white matter. These occur both subjacent to an injection site in area 17, and below labeled foci in area 18 projecting back to area 17, as if interstitial neurons shared the connectivity of overlying layer 6.

Animals