Cat area 17. II. Response properties of infragranular layer neurons in the absence of supragranular layer activity

1986 ◽  
Vol 56 (4) ◽  
pp. 1074-1087 ◽  
Author(s):  
H. D. Schwark ◽  
J. G. Malpeli ◽  
T. G. Weyand ◽  
C. Lee

Response properties of cells in the infragranular layers of cortical area 17 of the cat were examined in the absence of input from supragranular layers. Supragranular activity was silenced either reversibly by cooling the surface of cortex or permanently by making a cryogenic lesion of the supragranular layers. Visually driven responses of cells throughout the cortical column were recorded with a linear array of electrodes. Most infragranular layer cells continued to be visually responsive in the absence of supragranular layer input. These cells were similar to normal infragranular layer cells on measures of visual responsiveness, orientation selectivity, and direction selectivity. Special complex, but not standard complex, cells were absent in layer 5 when supragranular layers were destroyed. We found no evidence for a selective effect of removal of supragranular activity on the response properties of cells in layer 6. We propose that the intracolumnar projection from the supragranular layers drives the special complex cells of layer 5, but is not necessary for the visual driving of most other infragranular layer cells. This projection does not impose selectivity for stimulus orientation or direction on the remaining active cells of the infragranular layers.

1983 ◽  
Vol 49 (3) ◽  
pp. 595-610 ◽  
Author(s):  
J. G. Malpeli

1. Injections of 4 mM cobaltous chloride were used to block synaptic transmission in layer A of the lateral geniculate nucleus (LGN) without blocking fibers of passage going to or arising from other layers. 2. Selective inactivation of geniculate layer A virtually abolished all visual activity in cortical layers 4ab, 4c, and 6. Under these conditions, the stimulus-evoked response, orientation selectivity, and direction selectivity of cells in layers 2 and 3 were not seriously affected. In layer 5, the effects of the block were more variable, with special complex cells least affected and simple cells most affected. 3. Since the organization of complex receptive fields and the maintenance of normal orientation selectivity in supragranular layers survive disruption of major interlaminar interactions, it appears that much of the functional architecture of cat striate cortex does not depend on the integrity of the column. 4. These results support the idea that each layer of the LGN is a functional unit with a unique pattern of access to the various layers of visual cortex.


1976 ◽  
Vol 39 (3) ◽  
pp. 512-533 ◽  
Author(s):  
J. R. Wilson ◽  
S. M. Sherman

1. Receptive-field properties of 214 neurons from cat striate cortex were studied with particular emphasis on: a) classification, b) field size, c) orientation selectivity, d) direction selectivity, e) speed selectivity, and f) ocular dominance. We studied receptive fields located throughtout the visual field, including the monocular segment, to determine how receptivefield properties changed with eccentricity in the visual field.2. We classified 98 cells as "simple," 80 as "complex," 21 as "hypercomplex," and 15 in other categories. The proportion of complex cells relative to simple cells increased monotonically with receptive-field eccenticity.3. Direction selectivity and preferred orientation did not measurably change with eccentricity. Through most of the binocular segment, this was also true for ocular dominance; however, at the edge of the binocular segment, there were more fields dominated by the contralateral eye.4. Cells had larger receptive fields, less orientation selectivity, and higher preferred speeds with increasing eccentricity. However, these changes were considerably more pronounced for complex than for simple cells.5. These data suggest that simple and complex cells analyze different aspects of a visual stimulus, and we provide a hypothesis which suggests that simple cells analyze input typically from one (or a few) geniculate neurons, while complex cells receive input from a larger region of geniculate neurons. On average, this region is invariant with eccentricity and, due to a changing magnification factor, complex fields increase in size with eccentricity much more than do simple cells. For complex cells, computations of this geniculate region transformed to cortical space provide a cortical extent equal to the spread of pyramidal cell basal dendrites.


Perception ◽  
1997 ◽  
Vol 26 (1_suppl) ◽  
pp. 67-67
Author(s):  
S V Alexeenko ◽  
S N Toporova ◽  
F N Makarov

Intrinsic connections are one category of neuronal cortical connections, referring to axons that remain within the same cortical area and do not travel in the white matter. Our knowledge about these connections is based on extrapolations of results on the labelling of cells in different cortical layers, but there is no direct information about the complete connectivity of a single cortical column. To study how the population of cells sending axons to the same cortical column is distributed over the cortex we microiontophoretically injected horseradish peroxidase in single cortical columns of area 17 in cat. 3-D reconstruction of the region of labelled cells was performed by using serial frontal brain sections. It was shown that afferents from layer IV cells are short range (up to 0.5 mm) and from the supragranular (II, III) and the infragranular (V, VI) layer cells are long range (up to 5 mm). The regions with labelled cells in the supragranular and infragranular layers sending axons to the very same cortical column lie in register. For columns representing the central visual field, the distribution of labelling (in a tangential plane) is elongated in the mediolateral direction, and for peripheral columns there was a tendency towards elongation in the mediolateral and rostrocaudal directions. Moreover, the majority of labelled cells were located in regions representing peripheral parts of the visual field relative to the injection site. The described connections may represent the substrate for global linking tasks and underlie several psychophysical phenomena, such as meridional and peripheral effects.


1986 ◽  
Vol 56 (4) ◽  
pp. 1062-1073 ◽  
Author(s):  
J. G. Malpeli ◽  
C. Lee ◽  
H. D. Schwark ◽  
T. G. Weyand

Reversible inactivation of individual layers of the cat lateral geniculate and medial interlaminar nuclei was used to investigate the necessary and sufficient inputs for maintaining visually driven activity and receptive field properties in area 17. Neither orientation selectivity nor direction selectivity depends on any individual geniculate layer. We identified two groups of cortical layers on the basis of the pattern of thalamic inputs providing visual driving through the contralateral eye. One group, consisting of layers 4 and 6, has geniculate layer A as its only necessary and sufficient input. The other, consisting of supragranular layers, integrates at least two sufficient thalamic inputs, one of which is layer A. Several major receptive field properties are independently generated in these two groups of layers.


1987 ◽  
Vol 57 (5) ◽  
pp. 1511-1535 ◽  
Author(s):  
J. Cremieux ◽  
G. A. Orban ◽  
J. Duysens ◽  
B. Amblard

The response properties of 196 area 17 cells were studied qualitatively in seven cats reared from birth in a stroboscopically illuminated environment (frequency, 2/s; duration, 200 microseconds). Quantitative testing with the multihistogram technique was carried out in 115 cells. As control population, 453 neurons recorded in area 17 of the normal adult cat and tested qualitatively (of which 301 neurons were tested quantitatively) were available. In area 17 of strobe-reared cats, a number of spatial characteristics of receptive fields investigated with hand-held stimuli were found to be abnormal. There was a strong reduction in the encounter frequency both of end-stopped cells and of binocularly driven cells in the strobe-reared cats. Central receptive fields in strobe-reared cats were wider than in normal cats, but the increase in receptive-field width with eccentricity was still observed. More cells than in normal cats showed either no selectivity or only a weak bias for stimulus orientation, but the orientation tuning of orientation-selective cells was similar in strobe-reared and normal cats. Quantitative testing revealed that the velocity preference of cells in area 17 subserving central vision was different in strobe-reared cats from that of normal cats, due to a reduction in the encounter frequency of cells showing a preference for low velocities. There was no difference in velocity preference between strobe-reared and normal cats in the parts of area 17 that subserve peripheral vision, the proportion of neurons responding to fast velocities showing a similar increase in both groups of animals. Fewer cells were direction selective in strobe-reared cats than in normal cats. Most of the remaining direction-selective cells had peripheral receptive fields and the synergism between leaving an OFF subregion and entering an ON subregion contributed to their direction selectivity. Latency of neurons in area 17 of strobe-reared cats was slightly higher than in normal cats, but the response strength of neurons was the same in the two groups. The proportion of cells failing to respond to briefly flashed stationary stimuli was significantly lower in strobe-reared than in normal animals. Qualitative and quantitative testing showed that strobe rearing has a stronger effect on the parts of area 17 that subserve central vision than on those that subserve peripheral vision. Comparing the present results with those of Kennedy and Orban (37) shows that strobe rearing has less effect on area 17 than on area 18 and that the functional differences between areas 17 and 18 in strobe-reared cats are smaller than in normal cats.


2017 ◽  
Author(s):  
Marjena Popović ◽  
Andrea K. Stacy ◽  
Mihwa Kang ◽  
Roshan Nanu ◽  
Charlotte E. Oettgen ◽  
...  

AbstractMany sensory neural circuits exhibit response normalization, which occurs when the response of a neuron to a combination of multiple stimuli is less than the sum of the responses to the individual stimuli presented alone. In the visual cortex, normalization takes the forms of cross-orientation suppression and surround suppression. At the onset of visual experience, visual circuits are partially developed and exhibit some mature features such as orientation selectivity, but it is unknown whether cross-orientation suppression or surround suppression are present at the onset of visual experience or require visual experience for their emergence. We characterized the development of these properties and their dependence on visual experience in ferrets. Visual experience was varied across three conditions: typical rearing, dark rearing, and dark rearing with daily exposure to simple sinusoidal gratings (14-16 hours total). Cross-orientation suppression and surround suppression were noted in the earliest observations, and did not vary considerably with experience. We also observed evidence of continued maturation of receptive field properties in the second month of visual experience: substantial length summation was observed only in the oldest animals (postnatal day 90); evoked firing rates were greatly increased in older animals; and direction selectivity required experience, but declined slightly in older animals. These results constrain the space of possible circuit implementations of these features.Significance StatementThe development of the brain depends on both nature – factors that are independent of the experience of an individual animal – and nurture – factors that depend on experience. While orientation selectivity, one of the major response properties of neurons in visual cortex, is already present at the onset of visual experience, it is unknown if response properties that depend on interactions among multiple stimuli develop without experience. We find that the properties of crossorientation suppression and surround suppression are present at eye opening, and do not depend on visual experience. Our results are consistent with the idea that a majority of the basic properties of sensory neurons in primary visual cortex are derived independent of the experience of an individual animal.


1989 ◽  
Vol 3 (3) ◽  
pp. 249-265 ◽  
Author(s):  
Helen Sherk

AbstractThe existence of multiple areas of extrastriate visual cortex raises the question of how the response properties of each area are derived from its visual input. This question was investigated for one such area in the cat, referred to here as the Clare-Bishop area (Hubel & Wiesel, 1969); it is the region of lateral suprasylvian cortex that receives input from area 17. A novel approach was used, in which kainic acid was injected locally into the Clare-Bishop area, making it possible to record directly from afferent inputs.The response properties of the great majority of a sample of 424 presumed afferents resembled cells in areas 17 and 18. Thus, a systematic comparison was made with cells from area 17's upper layers, the source of its projection to the Clare-Bishop area (Gilbert & Kelly, 1975), to see whether these afferents had distinctive properties that might distinguish them from cells projecting to areas 18 or 19. Some differences did emerge: (1) The smallest receptive fields typical of area 17 were relatively scarce among afferents. (2) Direction-selective afferents were more abundant than were such cells in area 17. (3) End-stopped afferents were extremely rare, although end-stopped cells were common in area 17's upper layers.Despite these differences, afferents were far more similar in their properties to cells in areas 17 and 18 than to cells in the Clare-Bishop area. Compared to the latter, afferents showed major discrepancies in receptive-field size, in direction selectivity, in end-stopping, and in ocular dominance distribution. These differences seem most likely to stem from circuitry intrinsic to the Clare-Bishop area.


1990 ◽  
Vol 63 (6) ◽  
pp. 1529-1543 ◽  
Author(s):  
M. S. Gizzi ◽  
E. Katz ◽  
R. A. Schumer ◽  
J. A. Movshon

1. We consider the consequences of the orientation selectivity shown by most cortical neurons for the nature of the signals they can convey about the direction of stimulus movement. On theoretical grounds we distinguish component direction selectivity, in which cells are selective for the direction of movement of oriented components of a complex stimulus, from pattern direction selectivity, or selectivity for the overall direction of movement of a pattern irrespective of the directions of its components. We employed a novel test using grating and plaid targets to distinguish these forms of direction selectivity. 2. We studied the responses of 280 cells from the striate cortex and 107 cells from the lateral suprasylvian cortex (LS) to single sinusoidal gratings to determine their orientation preference and directional selectivity. We tested 73 of these with sinusoidal plaids, composed of two sinusoidal gratings at different orientations, to study the organization of the directional mechanisms within the receptive field. 3. When tested with single gratings, the directional tuning of 277 oriented cells in area 17 had a mean half width of 20.6 degrees, a mode near 13 degrees, and a range of 3.8-58 degrees. Simple cells were slightly more narrowly tuned than complex cells. The selectivity of LS neurons for the direction of moving gratings is not markedly different from that of neurons in area 17. The mean direction half width was 20.7 degrees. 4. We evaluated the directional selectivity of these neurons by comparing responses to stimuli moved in the optimal direction with those elicited by a stimulus moving in the opposite direction. In area 17 about two-thirds of the neurons responded less than half as well to the non-preferred direction as to the preferred direction; two-fifths of the units responded less than one-fifth as well. Complex cells showed a somewhat greater tendency to directional bias than simple cells. LS neurons tended to have stronger directional asymmetries in their response to moving gratings: 83% of LS neurons showed a significant directional asymmetry. 5. Neurons in both areas responded independently to each component of the plaid. Thus cells giving single-lobed directional-tuning curves to gratings showed bilobed plaid tuning curves, with each lobe corresponding to movement in an effective direction by one of the two component gratings within the plaid. The two best directions for the plaids were those at which one or other single grating would have produced an optimal response when presented alone.(ABSTRACT TRUNCATED AT 400 WORDS)


1991 ◽  
Vol 65 (5) ◽  
pp. 1078-1088 ◽  
Author(s):  
T. G. Weyand ◽  
J. G. Malpeli ◽  
C. Lee

1. We examined the response properties and geniculate inputs of 35 antidromically identified corticotectal (CT) cells within area 18 of the paralyzed, anesthetized cat. Twenty-three were either standard complex or hypercomplex, 11 were special complex, and 1 was simple. 2. The response properties of CT cells in area 18 were in general quite similar to those examined in a previous study of area 17 CT cells, including similar proportions of standard and special complex CT cells, virtually identical length-response functions, and similar orientation and direction tuning. 3. Area 18 CT cells are rapidly conducting. They are considerably faster than area 17 CT cells. 4. We investigated the composition of thalamic inputs to CT cells by reversibly inactivating a portion of layer A and/or the C layers of the dorsal lateral geniculate nucleus with injections of cobaltous chloride. Blocking layer A strongly attenuated the visual responsiveness of about half of the cells tested. Blocking the C layers alone generally had only moderate effects, but simultaneous blockade of layer A and the C layers demonstrated a substantial C-layer input to many cells. Unlike area 17 in which there is a strong correlation between CT cell class and dependence on layer A, no single receptive-field parameter nor set of parameters was correlated with dependence on layer A. However, cells least affected by simultaneous blockade of layer A and the C layers were special complex, suggesting that, as in area 17, area 18 special complex CT cells integrate more geniculate inputs than standard complex CT cells. 5. We propose that the similarities of response properties of area 17 and area 18 CT cells results from their participation in similar interlaminar columnar circuits and that differences in the patterns of geniculate control reflect differences in the global patterns of geniculate inputs to these two areas.


1986 ◽  
Vol 56 (4) ◽  
pp. 1088-1101 ◽  
Author(s):  
T. G. Weyand ◽  
J. G. Malpeli ◽  
C. Lee ◽  
H. D. Schwark

The receptive field properties of antidromically identified corticotectal (CT) cells in area 17 were explored in the paralyzed, anesthetized cat. To compare these with another population of infragranular cells, we also examined the receptive field properties of cells in layer 6. Sixty percent of our sample of CT cells showed increased response to increased stimulus length (length summation) and were classified as standard complex cells. The other 40% showed little or no length summation, were generally end stopped, and were classified as special complex cells. Standard and special complex CT cells have complementary orientation anisotropies: the distribution of orientation preferences of standard complex cells is biased toward obliquely oriented stimuli, whereas special complex cells are biased toward horizontally and vertically oriented stimuli. The receptive fields of the cells in our sample were primarily along the horizontal meridian so we cannot determine if these anisotropies are defined relative to the vertical meridian or relative to the meridian passing through the receptive field. The effects of these anisotropies in preferred orientation are minimized by the broad orientation tuning of CT cells. There was no simple relationship between the direction bias of CT cells and the reported direction bias of tectal cells. In contrast to the heterogeneity of corticotectal cells, layer 6 cells uniformly showed strong length summation, tight orientation tuning, and little spontaneous activity.


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