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PubMed1990

Temporal encoding of two-dimensional patterns by single units in primate primary visual cortex. I. Stimulus-response relations.

1. Previously we developed a new approach for investigating visual system neuronal activity in which single neurons are considered to be communication channels transmitting stimulus-dependent codes in their responses. Application of this approach to the stimulus-response relations of inferior temporal (IT) neurons showed that these carry stimulus-dependent information in the temporal modulation as well as in the strength of their responses. IT cortex is a late station in the visual processing stream. Presumably the neuronal properties arise from the properties of the inputs. However, the discovery that IT neuronal spike trains transmit information in stimulus-dependent temporally modulated codes could not be assumed to be true for those earlier stations, so the techniques used in the earlier study were applied to single-striate cortical neurons in the studies reported here. 2. Single-striate cortical neurons were recorded from three awake, fixating rhesus monkeys. The neurons were stimulated by two sets of patterns. The first set was made up of 128 black-and-white patterns based on a complete, orthogonal set of two-dimensional Walsh-Hadamard functions. These stimuli appear as combinations of black-and-white rectangles and squares, and they fully span the range of all possible black-and-white pictures that can be constructed in an 8 x 8 grid. Except for the stimulus that appeared as an all-white or all-black square, each stimulus had equal areas of white and black. The second stimulus set was made up of single bars constructed in the same 8 x 8 grid as the Walsh stimuli. These were presented both as black against a gray background and white against a gray background. The stimuli were centered on the receptive field, and each member of the stimulus set was presented once before any stimulus appeared again. 3. The responses of 21 striate cortical neurons were recorded and analyzed. Two were identified as simple cells and the other 19 as complex cells according to the criteria originally used by Hubel and Wiesel. The stimulus set elicited a wide variety of response strengths and patterns from each neuron. The responses from both the bars and the Walsh set could be used to differentiate and classify simple and complex cells. 4. The responses of both simple and complex cells showed striking stimulus-related strength and temporal modulation. For all of the complex cells there were instances where the responses to a stimulus and its contrast-reversed mate were substantially different in response strength or pattern, or both.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1990

Temporal encoding of two-dimensional patterns by single units in primate primary visual cortex. II. Information transmission.

1. Previously, we studied how picture information was processed by neurons in inferior temporal cortex. We found that responses varying in both response strength and temporal waveform carried information about briefly flashed stationary black-and-white patterns. Now, we have applied that same paradigm to the study of striate cortical neurons. 2. In this approach the responses to a set of basic black and white pictures were quantified through use of a set of basic waveforms, the principal components (extracted from all the responses of each neuron). We found that the first principal component, which corresponds to the response strength, and others, which correspond to different basic temporal activity patterns, were significantly related to the stimuli, i.e., the stimulus drove both the response strength and its temporal pattern. 3. Our previous study had shown that, when information theory was used to quantify the stimulus-response relation, inferior temporal neurons convey over twice as much information in a response code that includes temporal modulation as in a response code that includes only the response strength. This study shows that striate cortical neurons also carry twice as much information in a temporal code as in a response strength code. Thus single visual neurons at both ends of a cortical processing chain for visual pattern use a multidimensional temporal code to carry stimulus-related information. 4. These results support our multiplex-filter hypothesis, which states that single visual system neurons can be regarded as several simultaneously active parallel channels, each of which conveys independent information about the stimulus.

Analysis of Variance↗
PubMed1993

Visual resolution and sensitivity of single cells in the primary visual cortex (V1) of a nocturnal primate (bush baby): correlations with cortical layers and cytochrome oxidase patterns.

1. This study describes the response properties of V1 cortical cells in a nocturnal primate and examines the receptive field organization of these cells in relationship to anatomically defined layers and cytochrome oxidase (CO) rich blobs and CO poor interblob compartments. Visual resolution and contrast sensitivity are consistent with other physiological and behavioral measures in this species. Comparisons are made with response properties of the same zones in macaque monkey, as well as of area 17 of a distantly related species (cat) that also occupies a nocturnal niche. 2. The responses of single cells to drifting sinusoidal gratings were recorded in V1 (striate cortex) of anesthetized, paralyzed bush babies (Galago crassicaudatus). Cells tended to be grouped with respect to ocular dominance, orientation preference, and direction selectivity. There was a high proportion of monocularly driven cells as in macaque monkey. Only 6% of the cells were nonoriented. These were poorly tuned complex cells and bore no resemblance to nonoriented lateral geniculate nucleus (LGN)-like cells reported in layer IV of macaque monkeys. Unidirectional cells were most frequently encountered in cortical layers that receive input from the magnocellular layers of the LGN. 3. Cells were classified as simple (31%) or complex (69%) according to standard criteria. Simple cells were significantly more narrowly tuned than complex cells for both orientation and spatial frequency. Complex cells had significantly higher average optimal spatial frequencies and spatial frequency cutoffs than simple cells. Contrast sensitivity of simple and complex cells averaged 38 and 34, respectively. Spatial resolution and sensitivity of these cells matches behavioral measures in bush baby. The spatial and temporal resolution of bush baby cells are similar to those of cats, which is likely related to the nocturnal niche of both species. 4. Cells in supragranular (I-III) and infragranular (V, VI) layers differed significantly in their response characteristics. The cells in the supragranular layers had significantly higher contrast sensitivity than did the cells in the infragranular layers. Cells in the supragranular layers likewise had higher temporal frequency cutoffs, significantly lower optimal spatial frequencies, lower spatial frequency cutoffs, and tighter orientation tuning than did cells in the infragranular layers. 5. Properties of cells in individual layers and CO blob and interblob compartments also showed differentiation. Layer III had the narrowest orientation and spatial frequency tuning with the tightest tuning in layer IIIC (IVB).(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Length and width tuning of neurons in the cat's primary visual cortex.

1. The classically defined receptive field of a visual neuron is the area of visual space over which the cell responds to visual stimuli. It is well established, however, that the discharge produced by an optimal stimulus can be modulated by the presence of additional stimuli that by themselves do not produce any response. This study examines inhibitory influences that originate from areas located outside of the classical (i.e., excitatory) receptive field. Previous work has shown that for some cells the response to a properly oriented bar of light becomes attenuated when the bar extends beyond the receptive field, a phenomenon known as end-inhibition (or length tuning). Analogously, it has been shown that increasing the number of cycles of a drifting grating stimulus may also inhibit the firing of some cells, an effect known as side-inhibition (or width tuning). Very little information is available, however, about the relationship between end- and side-inhibition. We have examined the spatial organization and tuning characteristics of these inhibitory effects by recording extracellularly from single neurons in the cat's striate cortex (Area 17). 2. For each cortical neuron, length and width tuning curves were obtained with the use of rectangular patches of drifting sinusoidal gratings that have variable length and width. Results from 82 cells show that the strengths of end- and side-inhibition tend to be correlated. Most cells that exhibit clear end-inhibition also show a similar degree of side-inhibition. For these cells, the excitatory receptive field is surrounded on all sides by inhibitory zones. Some cells exhibit only end- or side-inhibition, but not both. Data for 28 binocular cells show that length and width tuning curves for the dominant and nondominant eyes tend to be closely matched. 3. We also measured tuning characteristics of end- and side-inhibition. To obtain these data, the excitatory receptive field was stimulated with a grating patch having optimal orientation, spatial frequency, and size, whereas the end- or side-inhibitory regions were stimulated with patches of gratings that had a variable parameter (such as orientation). Results show that end- and side-inhibition tend to be strongest at the orientation and spatial frequency that yield maximal excitation. However, orientation and spatial frequency tuning curves for inhibition are considerably broader than those for excitation, suggesting that inhibition is mediated by a pool of neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1999

Neuronal origin of the transcorneal electrically evoked response in area 17 (primary visual cortex) of cats.

We investigated how the neuronal activity of the cells in area 17 of cats evoked by transcorneal electrical stimuli is associated with the transcorneal electrically evoked response (EER). Neuronal firings were associated with the negative components of the EER, N1 (9 ms latency) and N2 (20 ms latency). Excitation of a longer latency was suppressed and periodic firings were rarely found. These findings indicate that the early negative components of the EER consist of the synaptic activities over the discharge thresholds of cortical cells, and also suggest that inhibition in the visual cortex may affect the EER.

Animals↗
PubMed1997

Topographic receptive fields and patterned lateral interaction in a self-organizing model of the primary visual cortex.

This article presents a self-organizing neural network model for the simultaneous and cooperative development of topographic receptive fields and lateral interactions in cortical maps. Both afferent and lateral connections adapt by the same Hebbian mechanism in a purely local and unsupervised learning process. Afferent input weights of each neuron self-organize into hill-shaped profiles, receptive fields organize topographically across the network, and unique lateral interaction profiles develop for each neuron. The model demonstrates how patterned lateral connections developed based on correlated activity and explains why lateral connection patterns closely follow receptive field properties such as ocular dominance.

Brain Mapping↗
PubMed1999

Development and organization of ocular dominance bands in primary visual cortex of the sable ferret.

Thalamocortical afferents in the visual cortex of the adult sable ferret are segregated into eye-specific ocular dominance bands. The development of ocular dominance bands was studied by transneuronal labeling of the visual cortices of ferret kits between the ages of postnatal day 28 (P28) and P81 after intravitreous injections of either tritiated proline or wheat germ agglutinin-horseradish peroxidase. Laminar specificity was evident in the youngest animals studied and was similar to that in the adult by P50. In P28 and P30 ferret kits, no modulation reminiscent of ocular dominance bands was detectable in the pattern of labeling along layer IV. By P37 a slight fluctuation in the density of labeling in layer IV was evident in serial reconstructions. By P50, the amplitude of modulation had increased considerably but the pattern of ocular dominance bands did not yet appear mature. The pattern and degree of modulation of the ocular dominance bands resembled that in adult animals by P63. Flat mounts of cortex and serial reconstructions of layer IV revealed an unusual arrangement of inputs serving the two eyes in the region rostral to the periodic ocular dominance bands. In this region, inputs serving the contralateral eye were commonly fused along a mediolateral axis, rostral to which were large and sometimes fused patches of ipsilateral input.

Aging↗
PubMed1999

Functional micro-organization of primary visual cortex: receptive field analysis of nearby neurons.

It is well established that multiple stimulus dimensions (e.g., orientation and spatial frequency) are mapped onto the surface of striate cortex. However, the detailed organization of neurons within a local region of striate cortex remains unclear. Within a vertical column, do all neurons have the same response selectivities? And if not, how do they most commonly differ and why? To address these questions, we recorded from nearby pairs of simple cells and made detailed spatiotemporal maps of their receptive fields. From these maps, we extracted and analyzed a variety of response metrics. Our results provide new insights into the local organization of striate cortex. First, we show that nearby neurons seldom have very similar receptive fields, when these fields are characterized in space and time. Thus, there may be less redundancy within a column than previously thought. Moreover, we show that correlated discharge increases with receptive field similarity; thus, the local dissimilarity between neurons may allow for noise reduction by response pooling. Second, we show that several response variables are clustered within striate cortex, including some that have not received much attention such as response latency and temporal frequency. We also demonstrate that other parameters are not clustered, including the spatial phase (or symmetry) of the receptive field. Third, we show that spatial phase is the single parameter that accounts for most of the difference between receptive fields of nearby neurons. We consider the implications of this local diversity of spatial phase for population coding and construction of higher-order receptive fields.

Animals↗
PubMed1990

Color and contrast sensitivity in the lateral geniculate body and primary visual cortex of the macaque monkey.

We tested color and contrast sensitivity in the magnocellular and parvocellular subdivisions of the lateral geniculate body and in layers 2, 3, 4B, and 4C alpha of visual area 1 to obtain physiological data on the degree of segregation of the 2 pathways and on the fate of the color and contrast information as it is transmitted from the geniculate to the cortex. On average, magnocellular geniculate cells were much less responsive than parvocellular cells to shifts between 2 equiluminant colors. Nevertheless, many magnocellular cells (though not all) continued to give some response at equiluminance. As expected from previous studies, luminance contrast sensitivity differed markedly between magnocellular and parvocellular layers. In V-1, the properties of cells in the magnorecipient layers 4C alpha and 4B faithfully reflected the properties of magnocellular geniculate cells, showing no evidence of any parvocellular input. Like magnocellular geniculate cells, they showed high contrast sensitivity, and with color contrast stimuli they showed large response decrements at equiluminance. In the interblob regions of cortical layers 2 and 3, which anatomically appear to receive most of their inputs from parvorecipient layer 4C beta, contrast sensitivities of some of the cells were compatible with a predominantly parvocellular input. Other interblob cells had sensitivities intermediate between magno- and parvocellular geniculate cells, suggesting a possible contribution from the magnocellular system. Many cells in cortical layers 2 and 3 responded to color-contrast borders equally well at all relative brightnesses of the 2 colors, including equiluminance. We recorded from many direction- and disparity-selective cells in V-1: most of the direction-selective and all of the clearly stereo-selective cells were located in layer 4B.

Animals↗
PubMed1986

Orientation selectivity of synaptic potentials in neurons of cat primary visual cortex.

Neurons of the visual cortex of the cat were penetrated with intracellular electrodes and postsynaptic potentials evoked by visual stimuli recorded. By alternately polarizing the cell with steady current injected through the recording electrode, IPSPs and EPSPs could be recorded and analyzed independently. Hyperpolarizing current suppressed IPSPs and enhanced EPSPs by moving the membrane potential toward the IPSP equilibrium potential. Depolarizing the cell toward the EPSP equilibrium potential enhanced IPSP. The responses to electrical stimulation of the LGN, where EPSPs and IPSPs could be distinguished easily by virtue of their characteristic latencies and shapes, were used to set the current injection to the appropriate level to view the two types of synaptic potential. EPSPs were found to be well oriented in that maximal depolarizing responses could be evoked at only one stimulus orientation; rotating the stimulus orientation in either direction produced a fall in the EPSP response. IPSPs were also well tuned to orientation, and invariably the preferred orientations of EPSPs and IPSPs in any one cell were identical. In addition, no systematic difference in the width of tuning of the two types of potential was seen. This result has been obtained from penetrations of over 30 cortical cells, including those with simple and complex receptive fields. It is concluded that orientation of cortical receptive fields is neither created nor sharpened by inhibition between neurons with different orientation preference. The function of inhibition evoked simultaneously with excitation by optimally oriented stimuli has yet to be determined, though it is likely to be the mechanism underlying other cortical receptive field properties, such as direction selectivity and end-stopping.

Animals↗
PubMed1984

Specificity of intrinsic connections in primate primary visual cortex.

Several recent studies have suggested a patchy system of intrinsic lateral connections in area 17 of the macaque monkey. To see whether this pattern bore any relationship to the cytochrome oxidase blobs we made multiple tiny injections of horseradish peroxidase into layers 2 and 3 of area 17, small enough so that some of the injections (or their cores) were entirely inside a single blob, or entirely outside. When the injection centers were entirely in blobs, the label in layers 2 and 3 was transported preferentially to nearby blobs, avoiding nonblob areas. When the injections were in nonblob areas, the label was found predominantly in surrounding nonblob areas, avoiding the blobs. Besides this lateral transport, label was also present in the layers below 2 and 3: the label in layers 4 and 6 was very restricted, occupying roughly the diameter of the injection core and presumably representing axons of cell bodies at the injection site; in layer 5 diffusely labeled patches observed the same blob/nonblob segregation seen above layer 4.

Animals↗
PubMed1984

[Demonstration of an effect of (3H)-L-proline on the number of dendritic spines on stellate cells of the primary visual cortex in Macaca during the critical period].

A morphometric study in cortical layer IVC of Macaca area 17 was performed during the critical period. A 20% increase in the number of dendritic spines was observed in neurons with horizontal dendritic tree in the cortical sublayer IVC alpha after an intraocular injection of (3H)-L-proline. A neuroactive role of this amino acid is discussed.

Animals↗