Nonlinearity of spatial summation in simple cells of areas 17 and 18 of cat visual cortex

1991 ◽  
Vol 66 (5) ◽  
pp. 1667-1679 ◽  
Author(s):  
D. Ferster ◽  
B. Jagadeesh
Keyword(s):  
Visual Cortex ◽  
Spatial Frequency ◽  
Spatial Summation ◽  
Area 17 ◽  
Simple Cells ◽  
Area 18 ◽  
Cat Visual Cortex ◽  
Spatial Frequencies ◽  
Areas 17 And 18 ◽  
Y Cells

1. Nonlinearity of spatial summation in areas 17 and 18 of cat visual cortex was compared with the type of spatial nonlinearity that differentiates X and Y cells in the lateral geniculate nucleus (LGN) and retina. The comparisons were made to examine to what extent the information from X and Y cells may remain separated in higher visual centers. 2. Responses of simple cells in areas 17 and 18 were recorded while stationary, optimally oriented sinewave gratings were sinusoidally modulated within the receptive field of the cell. Both the spatial frequency and spatial phase of the stimulus were varied. 3. Y cells in the retina and LGN are defined by the presence of a specific form of spatial nonlinearity. When tested with contrast-modulated sinewave gratings of spatial frequencies about three-fold greater than the optimal, their responses are dominated by a frequency-doubled component. The amplitude of the frequency-doubled component is not dependent on the spatial phase of the stimulus. 4. Many simple cells in the cortex showed a form of spatial nonlinearity similar to the defining nonlinearity found in retinal and geniculate Y cells. A frequency-doubled response dominated at spatial frequencies more than threefold greater than the optimal spatial frequency. When this response was present, it was phase independent. 5. More than 50% of the simple cells in area 18 showed the Y-like spatial nonlinearity. Fewer than 10% of the simple cells in area 17 showed the Y-like spatial nonlinearity. 6. The virtual absence of Y-like nonlinearity in area 17 and its relative abundance in area 18 suggest that the functional separation between the parallel X and Y pathways remains distinct within areas 17 and 18 of cat visual cortex.

On the variety of spatial frequency selectivities shown by neurons in area 17 of the cat

1981 ◽  
Vol 213 (1191) ◽  
pp. 183-199 ◽  
Keyword(s):  
Spatial Frequency ◽  
Visual Analysis ◽  
Frequency Tuning ◽  
Spatial Summation ◽  
Orientation Tuning ◽  
Area 17 ◽  
Tuning Curves ◽  
Spatial Frequencies ◽  
Functional Classes ◽  
Y Cells

The amplitudes of the responses of over 300 neurons in area 17 of the cat were examined as a function of the spatial frequency of moving sinusoidal gratings. The optimal spatial frequency and the bandwidth of the tuning curves were determined. The bandwidth varied considerably from neuron to neuron. Neurons optimally responsive to high spatial frequencies tended to have narrower tuning curves than those responsive to lower frequencies. Neurons with narrow spatial frequency tuning curves also tended to have narrow orientation tuning curves. These observations suggest that linear spatial summation tends to occur over a relatively constant area of visual field despite marked differences in each neuron’s optimal spatial frequency, a prediction of one model of visual analysis. There was little difference in either the optimal spatial frequencies or the bandwidths of tuning for different functional classes of neuron. Neurons with broad tuning curves tended to be restricted to lamina IV and its environs, being concentrated in the deep part of lamina II–III and the upper part of lamina IV ab. Neurons with very low optimal spatial frequencies were uncommon and tended to be found either at the border of laminae II–III and IV or in lamina V. These laminar distributions are discussed with respect to the laminar differences in the projection of l. g. n. X- and Y- cells to the visual cortex.

Effects of Feedback Projections From Area 18 Layers 2/3 to Area 17 Layers 2/3 in the Cat Visual Cortex

1999 ◽  
Vol 82 (5) ◽  
pp. 2667-2675 ◽  
Author(s):  
Susana Martinez-Conde ◽  
Javier Cudeiro ◽  
Kenneth L. Grieve ◽  
Rosa Rodriguez ◽  
Casto Rivadulla ◽  
...  
Keyword(s):  
Visual Cortex ◽  
Small Region ◽  
Orientation Tuning ◽  
Area 17 ◽  
Visual Responses ◽  
Area 18 ◽  
Cat Visual Cortex ◽  
Effects Of Feedback ◽  
Feedback Connections

In the absence of a direct geniculate input, area 17 cells in the cat are nevertheless able to respond to visual stimuli because of feedback connections from area 18. Anatomic studies have shown that, in the cat visual cortex, layer 5 of area 18 projects to layer 5 of area 17, and layers 2/3 of area 18 project to layers 2/3 of area 17. What is the specific role of these connections? Previous studies have examined the effect of area 18 layer 5 blockade on cells in area 17 layer 5. Here we examine whether the feedback connections from layers 2/3 of area 18 influence the orientation tuning and velocity tuning of cells in layers 2/3 of area 17. Experiments were carried out in anesthetized and paralyzed cats. We blocked reversibly a small region (300 μm radius) in layers 2/3 of area 18 by iontophoretic application of GABA and recorded simultaneously from cells in layers 2/3 of area 17 while stimulating with oriented sweeping bars. Area 17 cells showed either enhanced or suppressed visual responses to sweeping bars of various orientations and velocities during area 18 blockade. For most area 17 cells, orientation bandwidths remained unaltered, and we never observed visual responses during blockade that were absent completely in the preblockade condition. This suggests that area 18 layers 2/3 modulate visual responses in area 17 layers 2/3 without fundamentally altering their specificity.

Direction-sensitive X and Y cells within the A laminae of the cat's LGNd

1994 ◽  
Vol 11 (5) ◽  
pp. 927-938 ◽  
Author(s):  
Kirk G. Thompson ◽  
Yifeng Zhou ◽  
Audie G. Leventhal
Keyword(s):  
Visual Cortex ◽  
Spatial Frequency ◽  
Preferred Orientation ◽  
General Direction ◽  
Preferred Direction ◽  
X And Y Cells ◽  
Spatial Frequencies ◽  
Sinusoidal Gratings ◽  
Direction Sensitivity ◽  
Y Cells

AbstractDrifting sinusoidal gratings, moving bars, and moving spots were employed to study the direction sensitivity of 425 neurons in the A laminae of the cat's LGNd. Thirty-two percent of X- and Y-type LGNd relay cells exhibit significant direction sensitivity when tested with drifting sinusoidal gratings. X and Y cells exhibit the same degree of direction sensitivity. Moving spots and bars elicit direction specific responses from LGNd cells that are consistent with those elicited when drifting sinusoidal gratings are employed. For cells that are both orientation and direction sensitive, the preferred direction tends to be orthogonal to the preferred orientation. In general, direction sensitivity is strongest at relatively low spatial frequencies, well below the spatial-frequency cutoff for the cell. The presence of significant numbers of direction-sensitive LGNd cells raises the possibility that subcortical direction specificity is important for the generation of this property in the visual cortex.

Postnatal development of perisomatic GABAergic axon terminals on neurons projecting from area 17 to area 18 of the cat visual cortex

1999 ◽  
Vol 16 (1) ◽  
pp. 35-44 ◽  
Author(s):  
FERNANDO PÉREZ-CERDÁ ◽  
LUIS MARTÍNEZ-MILLÁN ◽  
CARLOS MATUTE
Keyword(s):  
Visual Cortex ◽  
Postnatal Development ◽  
Developmental Stages ◽  
Time Frame ◽  
Projection Neurons ◽  
Area 17 ◽  
Gamma Aminobutyric Acid ◽  
Area 18 ◽  
Axon Terminals ◽  
Cat Visual Cortex

We have studied the postnatal development of presumptive axon terminals (puncta) which were recognized by antibodies to the inhibitory neurotransmitter gamma-aminobutyric acid (GABA) and were located on the somata of area 17 neurons projecting to the ipsilateral area 18 of the visual cortex in cats ranging from 7 days of age to adulthood. Projection neurons were retrogradely labeled by injection of horseradish peroxidase conjugated to wheat germ agglutinin into the ipsilateral area 18. These neurons were mainly pyramidal in shape at all the developmental stages examined and the adult distribution of labeled cells was reached by 21 days. Subsequent GABA postembedding immunohistochemistry using high-resolution light microscopy was carried out to study the development of GABAergic terminals on cell bodies of identified projecting neurons in layers II–III. At all ages examined, we found perisomatic GABAergic puncta on these cells. Their density showed a significant increase from postnatal days 7 to 45, and then remained largely constant through adulthood. Since GABAergic puncta are considered the light-microscopic correlate of GABAergic synaptic terminals, our results support the idea of a developmentally regulated increase in the inhibitory activity of local interneurons on area 17 pyramidal neurons projecting to area 18 in the cat visual cortex which occurs within the same time frame as that of the acquisition of the mature operation of these cells.

Simple- and complex-cell response dependences on stimulation parameters

1985 ◽  
Vol 53 (5) ◽  
pp. 1244-1265 ◽  
Author(s):  
H. Spitzer ◽  
S. Hochstein
Keyword(s):  
Spatial Frequency ◽  
Time Course ◽  
Amplitude Dependence ◽  
Complex Cell ◽  
Simple Cells ◽  
Complex Cells ◽  
Stimulation Parameters ◽  
Spatial Frequencies ◽  
Y Cells ◽  
Two Peaks

We studied the response time course and amplitude dependence on stimulation parameters in cat cortical visual neurons to determine their receptive-field spatial-summation characteristics. Response poststimulus time (PST) histograms of cortical simple cells to contrast-reversal grating stimulation generally have a single peak for each stimulus temporal cycle, though the responses appear rectified. In response to contrast-reversal grating stimulation the general PST histogram time course for complex cells is two peaks, though often these peaks are of different amplitudes. The time course of complex-cell responses, and the ratio of these two response peaks often varies with stimulation parameters. The appearance of a single response peak in simple cells is reflected in the dominance of the odd harmonic Fourier portion, whereas the half-wave rectification leads to a considerable even harmonic portion. Still, this even portion is never significantly greater than the odd portion. When complex cell PST histograms have two nearly equal peaks, Fourier transformation reveals almost only even harmonic components. When the histogram contains two peaks of unequal amplitude Fourier analysis reveals large odd and even components. An even:odd Fourier harmonic portion ratio larger than 1 may be seen as a defining characteristic of complex cells, differentiating them from simple cells. Histograms with two unequal peaks appear "mixed," containing something of the "pure" single-peaked response and something of the pure double-peaked response. The degree to which the response is mixed may be measured by the ratio of the even:odd portion amplitudes. There is a great degree of variability with stimulation parameters (both spatial phase and spatial frequency) of the time course of mixed responses as opposed to the case of responses that have two equal peaks independent of stimulation grating phase and frequency. In both simple and complex cells there is a close coincidence of the spatial frequency ranges over which the even and odd portions are substantial, though many complex cells show a periodic variation of the even:odd portions ratio. This spatial-frequency dependence differs from that of LGN Y-cells where the odd portion dominates at low spatial frequencies and the even portion at high spatial frequencies. The ratio of even-to-odd portion cut-off is close to 3:1 in all Y-cells, a characteristic we did not find in cortical simple or complex cells. We suggest, therefore, that the nonlinearity of these complex cells does not derive from that of Y-cells.(ABSTRACT TRUNCATED AT 400 WORDS)

Stimulus dependence of orientation and direction sensitivity of cat LGNd relay cells without cortical inputs: A comparison with area 17 cells

1994 ◽  
Vol 11 (5) ◽  
pp. 939-951 ◽  
Author(s):  
Kirk G. Thompson ◽  
Audie G. Leventhal ◽  
Yifeng Zhou ◽  
Dan Liu
Keyword(s):  
Visual Cortex ◽  
Spatial Frequency ◽  
Frequency Tuning ◽  
Area 17 ◽  
Cortical Cells ◽  
Spatial Frequency Tuning ◽  
Spatial Frequencies ◽  
Direction Sensitivity ◽  
Cortical Inputs ◽  
Direction Tuning

AbstractThe cortical contribution to the orientation and direction sensitivity of LGNd relay cells was investigated by recording the responses of relay cells to drifting sinusoidal gratings of varying spatial frequencies, moving bars, and moving spots in cats in which the visual cortex (areas 17, 18, 19, and LS) was ablated. For comparison, the spatial-frequency dependence of orientation and direction tuning of striate cortical cells was investigated employing the same quantitative techniques used to test LGNd cells. There are no significant differences in the orientation and direction tuning to relay cells in the LGNd of normal and decorticate cats. The orientation and direction sensitivities of cortical cells are dependent on stimulus parameters in a fashion qualitatively similar to that of LGNd cells. The differences in the spatial-frequency bandwidths of LGNd cells and cortical cells may explain many of their differences in orientation and direction tuning. Although factors beyond narrowness of spatial-frequency tuning must exist to account for the much stronger orientation and direction preferences of cells in area 17 when compared to LGNd cells, the evidence suggests that the orientation and direction biases present in the afferents to the visual cortex may contribute to the orientation and direction selectivities found in cortical cells.

X- and Y-mediated synaptic potentials in neurons of areas 17 and 18 of cat visual cortex

1990 ◽  
Vol 4 (02) ◽  
pp. 115-133 ◽  
Author(s):  
David Ferster
Keyword(s):  
Visual Cortex ◽  
Optic Nerve ◽  
Cortical Neurons ◽  
Large Diameter ◽  
Area 18 ◽  
Synaptic Potentials ◽  
Areas 17 And 18 ◽  
Different Diameters ◽  
X Cells ◽  
Y Cells

AbstractWhen a cuff-shaped electrode is placed on the optic nerve of the cat, X and Y axons, by virtue of their different diameters, exhibit different thresholds to electrical stimulation. Large-diameter Y axons have low thresholds, while smaller-diameter X axons have high thresholds. There is very little overlap between the two populations. Given this segregation, the strength of stimulation of the optic nerve required to evoke synaptic potentials in cortical neurons becomes a reliable indicator of the type of visual input a cortical neuron receives. Potentials with thresholds below the thresholds of X axons must be mediated by Y cells of the retina and LGN. Potentials with thresholds above the Y axons of the optic nerve must be mediated by X cells. From previous experiments, one would expect ot find &le input via both types of axon to are 17 of the visual cortex. This was not the case. Of 58 neurons distributed throughout the layers of area 17 from which intracellular records were taken, in only four could substantial Y excitation be detected. Three of these four were located near the border with area 18. All four received large X inputs as well. The 24 neurons studied in area 18 all received large Y inputs but no detectable X input.

Visual callosal projections in the adult ferret

1992 ◽  
Vol 9 (1) ◽  
pp. 99-103 ◽  
Author(s):  
Antony M. Grigonis ◽  
Rosemary B. Rayos Del Sol-Padua ◽  
E. Hazel Murphy
Keyword(s):  
Visual Cortex ◽  
Horseradish Peroxidase ◽  
Retinal Ganglion Cells ◽  
Ganglion Cells ◽  
Retinal Ganglion ◽  
Cell Distribution ◽  
Area 17 ◽  
Area 18 ◽  
Areas 17 And 18 ◽  
Radial Organization

AbstractThe laminar and tangential organization of visual callosal projections of areas 17 and 18 were investigated in the adult ferret, using histochemical methods to visualize axonally transported horseradish peroxidase (HRP). Normal adult ferrets were given injections of HRP throughout one visual cortex or had gelfoam soaked in HRP applied to the transected corpus callosum. The ferret callosal cell distribution has a greater tangential extent in area 18 than in area 17. In addition, the radial organization of callosal cells in areas 17 and 18 differs: three times as many infragranular cells are present in area 18 than in area 17, although the number of supragranular cells is similar for both areas 17 and 18. Since the projections of alpha retinal ganglion cells are reported to be exclusively contralateral in the ferret (Vitek et al., 1985), callosal projections may make a major contribution to the binocularity of neurons in area 18.

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