Binocular interaction fields of single units in the cat striate cortex

1. Based on average response histograms to an optimal stimulus, binocular interaction field plots were obtained from twenty-five simple neurones in the striate cortex of the cat. Each binocularly activated cell has two interaction fields, one for each eye. The binocular interaction field for one eye plots the changes in the amplitude of the response from the other eye as the two receptive fields of the binocularly activated cell are moved across one another, first into and then out of alignment in the plane of the optimal stimulus (tangent screen).2. The binocular interaction field provides an important clue to the nature of the spatial organization of the excitatory and inhibitory regions of the monocular receptive field. The commonest type of receptive field organization has regions of inhibition (inhibitory side bands) to either side of the discharge centre in the direction at right angles to the optimal stimulus orientation. As well as inhibition, there are subliminal excitatory effects.3. Binocular interaction fields differ with the various cell types, i.e. cells that are discharged only from the one eye, cells binocularly discharged with very weak or absent monocular responses and cells showing binocularly opposite direction selectivity.4. Marked facilitation to an optimal stimulus occurs when the two receptive fields of a binocularly activated neurone are in accurate alignment. Facilitation switches to depression for very small degrees of receptive field misalignment in a direction at right angles to the optimal stimulus orientation. These observations are of importance in relation to mechanisms for binocular single vision and depth discrimination.     

Cutaneous masking. II. Geometry of excitatory andinhibitory receptive fields of single units in somatosensory cortex of the cat.

1. The responses of single neurons in the primary somatosensory cortex of the cat to brief air-pulse stimuli were quantitatively examined. These controlled natural stimuli activated almost exclusively rapidly adapting hair units which, on systematic movement of the stimulus through the receptive field, gave unit-response profiles that showed the classical unimodal tent-shaped distribution. 2. Conditioning stimulus-induced inhibition of a response evoked by a fixed test stimulus was measured by systematically moving the conditioning stimulus through the receptive field. The spatial distribution of in-field inhibitory activity was unimodal and highly covariant with that of the conditioning excitation, the peak inhibition corresponding to the functional center of the excitatory receptive field. 3. Nearly one-half of the units studied evidenced inhibition extending beyond the excitatory receptive field, forming a "surround" inhibitory region; but these were usually restricted areas with rather weak inhibitory effects. 4. Time-course measuring revealed, on the average, inhibition effects measureable from 10 ms before to some 70 ms following conditioning stimulation, with peak inhibition delayed some 10--15 ms from the conditioning stimulus onset. We showed the backward inhibition, occurring with the test stimulus delivered before the onset of the conditioning stimulus, to be a property of the test response duration. Inhibition measured in the surround areas had essentially the same time course as the inhibition calculated from measurements made within the receptive fields. 5. The spatial and temporal profiles of the excitatory and inhibitory cortical unitary activity are thus very similar to the parametric features of psychophysical enhancement and masking. These findings suggest that the excitatory and inhibitory activities related to individual stimuli interact in multipoint stimulus paradigms so that simple unimodal composite profiles are synthesized.     

Orientation selectivity and the spatial distribution of enhancement and suppression in receptive fields of cat striate cortex cells

The relationship between orientation selectivity and spatial receptive field organization was analyzed. Receptive field maps were made with a dual stimulus technique where an optimally oriented activation slit was presented in the most responsive region to produce activity against which the effect of a test spot in various positions was determined. Both simple and complex cells had receptive fields which were subdivided into adjacent elongated and antagonistic subregions. When the two stimuli were presented in phase (both ON or OFF simultaneously) the fields had a central enhancement region with a strong suppression flank on one or both sides. Optimal slit orientation was related to the location of the suppression flank relative to the location of the central enhancement region, and the degree of orientation selectivity to the shape of the subregions and the distance between them. Estimated orientation tuning curves calculated from the receptive field maps gave satisfactory first approximations to experimental curves. The relative contribution of enhancement and suppression to orientation selectivity was studied by presenting a test slit in different orientations in phase with an optimally oriented activation slit. The orientation selectivity was produced almost exclusively by the flank suppression indicating that orientation selectivity is produced by inhibitory input. The flank suppression lacked any specific orientation selectivity, and it occurred only when both the central region and the flanks were activated in phase. Orientation selectivity in both simple and complex cells is explained by a receptive field organization where the cells have input from partially overlapping excitatory and inhibitory fields which have their centers slightly displaced from each other.     

Responses of single units in the monkey superior colliculus to stationary flashing stimuli

1. Responses of pan-directional cells in the superficial layers of the superior colliculus in paralysed anaesthetized rhesus monkeys to stationary flashing stimuli have been studied. 2. The receptive field centre response is always of the transient excitatory on-off type, while the surround response is transient inhibitory both at light-on and at light-off. The receptive field centres are circular or slightly elliptical. The average size of the receptive field centres is much larger than that of retinal ganglion cells. All units except those in the far temporal periphery receive binocular input. In each unit the on and off responses have the same latency times. With increasing stimulus area, the latency time at light-on and at light-off first decreases and then remains constant. In most units the number of spikes in the burst at light-on and at light-off first increases, reaches a maximum and then decreases with increasing stimulus area. This decrease demonstrates the presence of an inhibitory surround. 3. A model of spatial and temporal properties of centre and surround mechanisms is tested. Addition of excitatory centre input and inhibitory surround input, which have different spatial and temporal properties, determines the output of the neurone. The centre mechanism gets excitatory input from retinal ganglion cells and shows saturation. The inhibitory surround mechanism is made by an inhibitory interneurone. It could not be decided whether the excitatory input for this interneurone comes from retinal axon collaterals (forward inhibition) or from axon collaterals of "principal" cells in the superior colliculus (backward inhibition).     

The properties of the binocular receptive fields of lateral geniculate neurons

Summary The majority of cells in the dorsal nucleus of the lateral geniculate body (LGNd) in the cat have two receptive fields: one for each eye. Of the cells tested for binocularity (113), only 21 (18%) were purely monocular. The remainder had receptive fields for the non-dominant eye, the great majority of which (81 or 88%) were purely inhibitory and only 11 (12%) were excitatory. Cells with receptive fields for the non-dominant eye were found in all three laminae (A, A₁ and B) of the LGNd. The proportion of inhibitory receptive fields for the non-dominant eye was slightly greater when the dominant eye was ipsilateral (77%) than when it was contralateral (68%). The distribution of the binocular receptive field pairs about points of exact correspondence in the visual field had a standard deviation of about 0.9° in both horizontal and vertical directions.The properties of the inhibitory receptive fields were studied with moving slits of light and stationary flashing spots. Most of the fields were purely inhibitory and varied in size from 1.5° to 6° across. There were no specific stimulus requirements other than a change in contrast within the receptive field. The inhibitory effect was usually fairly weak, the spontaneous discharge of the neuron being inhibited much more readily than the driven discharge. The latency of the inhibition to a stationary flashing spot was about 50 msec, the inhibition was maximal about 20 msec after the onset and lasted up to about 400 msec.Binocular inhibition is not mediated by a corticogeniculate pathway from the visual areas since it survives removal of areas 17, 18 and 19 and the middle suprasylvian gyrus. It was concluded that the most likely mechanism was via interneurons whose axons cross the borders from one cell layer to another.     

Responses to visual contours: spatio-temporal aspects of excitation in the receptive fields of simple striate neurones

1. The properties of the receptive fields of simple cells in the cat striate cortex have been studied by preparing average response histograms both to moving slits of light of different width and to single light-dark edges or contours.2. The movement of a narrow (< 0.3 degrees ) slit across the receptive field gives rise to average response histograms that are either unimodal, bimodal or multimodal. A slit of light has leading (light) and trailing (dark) edges. By increasing the width of the slit it was shown that a discharge peak in the histogram coincides with the passage of one or other of the two edges over a particular region (discharge centre) in the receptive field. Each edge has its own discharge centre which is fired when the edge has the correct orientation and direction of movement.3. The discharge centres in forty-three simple cell receptive fields were located by using one or more of the following stimuli for each cell:(i) slits of different width;(ii) single light and dark edges;(iii) a wide (3 degrees ) slit moved over a range of different velocities.The same locations were obtained when all three procedures were used on the same cell.4. Most cells (79%) discharged to both edges though not necessarily in the same direction of movement. The majority (72%) fired in only one direction and most commonly (51%) the cells responded to both edges in this one direction. In only 16% of cells did both types of edge excite in both directions of movement. When the one type of edge, light or dark, was considered, 84% of the cells were direction selective and, for these cells, the other edge fired only in the same direction (51%), in both directions (7%), only in the opposite direction (5%) or not at all (21%).5. Cells responding in one direction with a unimodal average response histogram may be responding to both edges, the two responses being concealed in the one discharge peak. The two discharge centres are then either nearly coincident or, more usually, slightly offset with respect to one another. Most commonly the dark edge centre is slightly in advance of the light edge centre.6. The discharge peaks in the bimodal and multimodal types come from discharge centres that are spatially separate, each centre firing to only one type of edge. In the case of the bimodal type the light edge centre always lies ahead of the dark edge centre.7. When a cell responds to a single edge in both directions of movement, the type of contrast effective in one direction is always the reverse of that in the other. When the cell responded in both directions, whether to one or both edges, most commonly a light edge discharge centre in one direction occupied approximately the same location in space as the dark edge centre in the reverse direction and vice versa for the other edge.8. Temporal aspects of the discharge of simple cells have been examined by recording the responses to moving slits and single edges over a wide range of velocities.     

Receptive field organization of complex cells in cat striate cortex

Summary The receptive field organization of complex cells was studied by analyzing interaction effects between two stationary flashing light stimuli. One was placed in the most responsive part of the receptive field to produce activity against which effects of the other in different visual field positions could be determined.The receptive field was spatially organized into antagonistic center and flanks just like the fields of simple cells. However, both center and flanks were found within the receptive field area where a single slit evoked discharge. Center and flanks were elongated along the optimal stimulus orientation. The flanks were displaced from the center normal to optimal stimulus orientation.In the center, ON- and OFF-responses were usually about equal in strength and the maximum ON- and OFF-responses occurred in about the same position. This shows that complex cells are activated by input from both ON- and OFF-center cells in the lateral geniculate nucleus (LGN) where the receptive field centers of the LGN cells overlap closely. This explains most of the specific features of complex cells, e.g., the spatially overlapping ON- and OFF-zones, the large response field, the repetitive firing when a slit moves over the receptive field, and the marked non-linear spatial summation.Strong flank suppression occurred with both ON and OFF. The effects were usually stronger on one side of the center. Maximal suppression occurred on the same side with both ON and OFF. This is consistent with the interpretation that complex cells are inhibited by input from both LGN ON- and OFF-center cells with overlapping receptive field centers.A model presuming that complex cells have overlapping but acentric excitatory and inhibitory fields was tested by computer simulation and shown to fit the experimental data. This is the same model as presented for simple cells in the preceding paper (Heggelund 1980), except that the excitatory and inhibitory fields of simple cells have input from either ON- or OFF-center LGN cells, whereas in complex cells they have input from both types.The project was financially supported by the Norwegian Research Council for Science and Humanities     

Single unit responses in the inferior colliculus of the awake and performing rhesus monkey

1. The activity of single units in the inferior colliculus of unanesthetized monkeys was recorded during performance in an auditory reaction time task. Stimulus intensity and frequency were varied. 2. Spontaneous rate of unit discharge varied from 0 to 78.2 discharges per second, with a mean of 14.7 discharges/sec. 3. Both broadly and narrowly tuned units were encountered in the central nucleus of the inferior colliculus. The temporal discharge pattern of most units varied with changes in stimulus frequency; onset bursts and/or sustained discharge suppression dominated the unit discharge at the edges of receptive fields. 4. Half of the units examined at several intensity levels displayed nonmonotonic relationships between evoked discharge rate and stimulus intensity, with most nonmonotonic units showing a distinct "best intensity". The temporal response pattern of almost all units varied with changes in stimulus intensity, with onset bursts and discharge suppression increasing in occurrence with increasing intensity. 5. Units recorded in the external nucleus of the inferior colliculus displayed spontaneous rates which were similar to those of central nucleus units, and were affected by variation in stimulus intensity in the same fashion. However, the average initial latency of such units to intense stimuli was no longer than the latency of central nucleus units. 6. Variations in unit discharge with changes in stimulus frequency and intensity are consistent with an interaction of excitatory and inhibitory inputs with different initial latencies, dynamic ranges and receptive fields. In particular, our data suggest that inhibitory inputs have longer initial latencies and higher thresholds. Inhibition is stronger at the edges of a unit's receptive field, and dominates at high frequencies in units with low characteristic frequency. 7. Our data are not consistent with previous reports that single units in the unanesthetized animal display uniformly monotonic intensity functions and uniformly broad frequency responses.     

Größenschätzung des rezeptiven Feldzentrums der menschlichen Retina

The antagonistic receptive field organization found in the vertebrate eye, caused by interplay of excitatory and inhibitory influences over interconnections within the retina, gives rise to a neural activity dependent on the shape of the spatial stimulus pattern falling onto the receptor mosaic. This phenomenon shows strong correlation with subjective contour-perception effects in human vision, and thus it is plausible that the same kinds of retinal mechanisms are prevalent in man. Assuming this to be so, the horizontal diameter of the foveal receptive field centres has been determined from measurements of the perceived luminance enhancement and reduction of the maximum- and minimum points, respectively, in a viewed one-dimensional sinusoidal spatially-varying stimulus pattern as a function of its spatial frequency and objective contrast. For object contrasts above 0.3 the diameter of the foveal receptive field centre was found to be about 28 μ. Moreover, the size of the receptive field centres belonging to on- as well as off-centre neurons were found to possess the same order of magnitude throughout this object contrast region. For lower object contrast values, on the other hand, response behaviour was found which seems to change the above mentioned size of the receptive field centres differently for on- and off-elements respectively.     

Gamma-aminobutyric acid and afferent inhibition in the cat and rat ventrobasal thalamus

Extracellular single neuron recordings were made in the ventrobasal thalami of anaesthetized rats and cats. Physiological stimulation of vibrissa and hair follicle afferents was performed with an air jet (10-20 ms duration) directed at a single vibrissa or small area of hairy skin. Paired conditioning and test air jets delivered to the excitatory portion of receptive fields revealed inhibition of the response of ventrobasal thalamic neurons to test stimuli following the excitatory response to the conditioning stimulus. Such inhibitions could last up to 500 ms. An increase in neuronal excitability was sometimes observed following this inhibitory period. In addition, it was possible to produce inhibition without an excitatory response using conditioning stimuli delivered adjacent to the excitatory receptive field. Iontophoretic application of bicuculline methochloride, with currents that were adequate to antagonize iontophoretically applied GABA, was found to reduce the inhibition of test responses evoked by conditioning stimuli in almost all of the neurons studied. In most cases, no excitatory responses to conditioning stimuli directed outside the original excitatory receptive field were revealed by application of the GABA antagonist. In rats, bicuculline also led to a decrease in the post-inhibitory excitation, whereas in cats the converse appeared to be the case. These results suggest that GABAergic transmission may underlie inhibitory responses of cat and rat ventrobasal thalamus neurons to physiological stimulation of somatosensory afferents. Furthermore, removal of such inhibition does not appear to reveal excitatory inputs from outside of the original excitatory receptive field.     

The spatial extent of excitatory and inhibitory zones in the receptive field of superficial layer hypercomplex cells

1. An investigation has been made of the extent of inhibitory and excitatory components in the receptive field of superficial layer hypercomplex cells in the cat's striate cortex and the relation of the components to the length preference exhibited by these cells.2. Maximal responses were produced by an optimal length stimulus moving through a restricted region of the receptive field. The length of this receptive field region was less than the total length of the excitatory zone as mapped with a very short slit. Slits of similar length to the excitatory zone produced a smaller response than an optimal length slit.3. An increase of slit length so that it passed over receptive field regions either side of the excitatory zone resulted in an elimination of the response. When background discharge levels were increased by the iontophoretic application of D, L-homocysteic acid slits of this length were observed to produce a suppression of the resting discharge as they passed over the receptive field. They did not modify the resting discharge level when it was induced by the iontophoretic application of the GABA antagonist bicuculline. This data is taken to indicate that long slits activate a powerful post-synaptic inhibitory input to the cell.4. Maximal inhibitory effects were only observed if the testing slit passed over the receptive field centre. That is slits with a gap positioned midway along their length so as to exclude the optimal excitatory response region surprisingly tended to produce excitatory effects rather than the expected inhibitory effects. It appears that simultaneous stimulation of the receptive field centre is a precondition for the inhibitory effect of stimulation of regions either side of the excitatory zone to be activated.5. It is suggested that the interneurones mediating the inhibitory input to the superficial layer hypercomplex cells are driven both by cells in adjacent hypercolumns with receptive fields spatially displaced to either side of the excitatory zone and by cells in the same column, optimal inhibitory effects only being achieved when both sets of input to the interneurone are activated.     

Dynamic properties of recurrent inhibition in primary visual cortex: contrast and orientation dependence of contextual effects

A fundamental feature of neural circuitry in the primary visual cortex (V1) is the existence of recurrent excitatory connections between spiny neurons, recurrent inhibitory connections between smooth neurons, and local connections between excitatory and inhibitory neurons. We modeled the dynamic behavior of intermixed excitatory and inhibitory populations of cells in V1 that receive input from the classical receptive field (the receptive field center) through feedforward thalamocortical afferents, as well as input from outside the classical receptive field (the receptive field surround) via long-range intracortical connections. A counterintuitive result is that the response of oriented cells can be facilitated beyond optimal levels when the surround stimulus is cross-oriented with respect to the center and suppressed when the surround stimulus is iso-oriented. This effect is primarily due to changes in recurrent inhibition within a local circuit. Cross-oriented surround stimulation leads to a reduction of presynaptic inhibition and a supraoptimal response, whereas iso-oriented surround stimulation has the opposite effect. This mechanism is used to explain the orientation and contrast dependence of contextual interactions in primary visual cortex: responses to a center stimulus can be both strongly suppressed and supraoptimally facilitated as a function of surround orientation, and these effects diminish as stimulus contrast decreases.     

Neural mechanisms for processing binocular information II. Complex cells.

Complex cells in the striate cortex exhibit extensive spatiotemporal nonlinearities, presumably due to a convergence of various subunits. Because these subunits essentially determine many aspects of a complex cell receptive field (RF), such as tuning for orientation, spatial frequency, and binocular disparity, examination of the RF properties of subunits is important for understanding functional roles of complex cells. Although monocular aspects of these subunits have been studied, little is known about their binocular properties. Using a sophisticated RF mapping technique that employs binary m-sequences, we have examined binocular interactions exhibited by complex cells in the cat's striate cortex and the binocular RF properties of their underlying functional subunits. We find that binocular interaction RFs of complex cells exhibit subregions that are elongated along the frontoparallel axis at different binocular disparities. Therefore responses of complex cells are largely independent of monocular stimulus position or phase as long as the binocular disparity of the stimulus is kept constant. The binocular interaction RF is well described by a sum of binocular interaction RFs of underlying functional subunits, which exhibit simple cell-like RFs and a preference for different monocular phases but the same binocular disparity. For more than half of the complex cells examined, subunits of each cell are consistent with the characteristics specified by an energy model, with respect to the number of subunits as well as relationships between the subunit properties. Subunits exhibit RF binocular disparities that are largely consistent with a phase mechanism for encoding binocular disparity. These results indicate that binocular interactions of complex cells are derived from simple cell-like subunits, which exhibit multiplicative binocular interactions. Therefore binocular interactions of complex cells are also multiplicative. This suggests that complex cells compute something analogous to an interocular cross-correlation of images for a local region of visual space. The result of this computation can be used for solving the stereo correspondence problem.     

Dynamic response properties of visual neurons and context-dependent surround effects on receptive fields in the tectum of the salamander Plethodon shermani

Neuronal responses to complex prey-like stimuli and rectangles were investigated in the tectum of the salamander Plethodon shermani using extracellular single-cell recording. Cricket dummies differing in size, contrast or movement pattern or a rectangle were moved singly through the excitatory receptive field of a neuron. Paired presentations were performed, in which a reference stimulus was moved inside and the different cricket dummies or the rectangle outside the excitatory receptive field. Visual object recognition involves much more complex spatial and temporal processing than previously assumed in amphibians. This concerns significant changes in absolute number of spikes, temporal discharge pattern, and receptive field size. At single presentation of stimuli, the number of discharges was significantly changed compared with the reference stimulus, and in the majority of neurons the temporal pattern of discharges was changed in addition. At paired presentation of stimuli, neurons mainly revealed a significant decrease in average spike number and a reduction of excitatory receptive field size to presentation of the reference stimulus inside the excitatory receptive field, when a large-sized cricket stimulus or the rectangle was located outside the excitatory receptive field. This inhibition was significantly greater for the large-sized cricket stimulus than for the rectangle, and indicates the biological relevance of the prey-like stimulus in object selection. The response properties of tectal neurons at single or paired presentation of stimuli indicate that tectal neurons integrate information across a much larger part of visual space than covered by the excitatory receptive field. The spike number of a tectal neuron and the spatio-temporal extent of its excitatory receptive field are not fixed but depend on the context, i.e. the stimulus type and combination. This dynamic processing corresponds with the selection of the stimuli in the visual orienting behavior of Plethodon investigated in a previous study, and we assume that tectal processing is modulated by top down processes as well as feedback circuitries.     

The contribution of inhibitory mechanisms to the receptive field properties of neurones in the striate cortex of the cat.

1. The iontophoretic application of the GABA antagonist bicuculline to simple and complex cells in the striate cortex of the cat produced extensive modifications of receptive field properties. These modifications appear to relate to a block or reduction of GABA-mediated intracortical inhibitory influences acting on the cells examined. 2. For simple cells the effects of bicuculline on receptive field properties involved a loss of the subdivision of the receptive field into antagonistic "on" and "off" regions, a reduction in orientation specificity and a reduction or elimination of directional specificity. 3. The effect on the "on" and "off" subdivisions of the simple cell receptive field was such that all stationary flashing stimuli, whether covering the whole receptive field, or located within the receptive field over a previously determined "on" or "off" region, resulted in an "on and off" response. 4. The orientation specificity of complex cells was reduced during the application of bicuculline such that in many cases the original specificity of the cell was virtually lost with the response to the orientation at 90 degrees to the optimal being of similar magnitude to the optimal. The directional specificity of complex cells was generally less affected than that of simple cells. Often when large changes in orientation specificity were observed the directional specificity was relatively unaffected. 5. For some cells apparently showing to all visual stimuli only inhibitory responses, the application of bicuculline resulted in the appearance of excitatory responses. 6. In all cases receptive field properties reverted to the original state after termination of the bicuculline application. It was not generally possible to duplicate the effects of bicuculline by raising neuronal excitability with iontophoretically applied glutamate. 7. On the basis of these results it is suggested that the normal subdivision of the simple cell receptive field into separate "on" and "off" regions and its directional specificity are dependent on intracortical inhibitory processes that are blocked by bicuculline. The orientational tuning of simple cells conversely appears to be largely determined by the excitatory input but normally enhanced by lateral type inhibitory processes acting in the orientation domain. 8. It also appears that the excitatory input to some complex cells is not orientation specific. This suggests that for these cells it is extremely unlikely that they receive an orientation specific excitatory input from simple cells.     

The binocular organization of complex cells in the cat's visual cortex

We have studied the manner by which inputs from the two eyes are combined in complex cells of the cat's visual cortex. The stimuli are drifting sinusoidal gratings presented dichoptically at optimal spatial frequency and orientation. The relative phase between the gratings for left and right eyes is varied over 360 degrees. Approximately 40% of complex cells show phase-specific binocular interaction where response amplitudes vary depending on the relative phase of the gratings shown to the two eyes. This interaction is similar to that observed for most simple cells. We devised a test to examine whether the phase-specific interaction in complex cells results from linear convergence of neural signals at subunits of the receptive fields. The data from this test are consistent with a linear combination model. The phase-specific binocular interaction data from complex cells imply that the optimal relative phase of the receptive field subunits is closely matched. Another type of complex cell, approximately 40% of the total, could be driven through either eye, but exhibited non-phase-specific responses to dichoptically presented gratings. This type of interaction is found only in complex cells. Binocularly non-phase-specific complex cells may have subunits whose optimal relative phases are random or monocular. The division of complex cells into these two major groups (binocularly phase specific and non-phase specific) is independent of whether they are standard or special complex-cell types. A small proportion (8%) of complex cells that appear monocular by alternate tests of each eye show a purely inhibitory influence from the silent eye. This inhibition is not generally dependent on the relative phase of the gratings. Unlike simple cells, complex cells are not a homogeneous group. However, nearly half of complex cells show phase-specific binocular interaction that is probably the result of linear convergence. Combined with the results from simple cells, the majority of binocular interaction in the striate cortex may be accounted for by linear summation of neural signals from each eye. This provides a simplified view of the nature of binocular interaction in the visual cortex.     

Properties of the receptive fields of frog retinal ganglion cells as revealed by their response to moving stimuli

Quantitative mapping of the excitatory receptive fields of frog retinal ganglion cells revealed that the precise spatial position and locus of maximal activity within the excitatory receptive field were dependent upon the exact sequence with which the moving stimulus scanned the region of visual space in which the excitatory receptive field was embedded. A first-order asymmetry dependent upon the direction from which the moving stimulus entered the excitatory receptive field was noted. A second-order asymmetry, orthogonal to the first and sensitive to the direction in which a moving stimulus systematically traversed successive columns with in the excitatory receptive field was also described. These findings indicate that complicated interactions, both excitatory and inhibitory, occur between component parts of the receptive field in the frog retina.     

Eye position compensation improves estimates of response magnitude and receptive field geometry in alert monkeys

Studies of visual function in behaving subjects require that stimuli be positioned reliably on the retina in the presence of eye movements. Fixational eye movements scatter stimuli about the retina, inflating estimates of receptive field dimensions, reducing estimates of peak responses, and blurring maps of receptive field subregions. Scleral search coils are frequently used to measure eye position, but their utility for correcting the effects of fixational eye movements on receptive field maps has been questioned. Using eye coils sutured to the sclera and preamplifiers configured to minimize cable artifacts, we reexamined this issue in two rhesus monkeys. During repeated fixation trials, the eye position signal was used to adjust the stimulus position, compensating for eye movements and correcting the stimulus position to place it at the desired location on the retina. Estimates of response magnitudes and receptive field characteristics in V1 and in LGN were obtained in both compensated and uncompensated conditions. Receptive fields were narrower, with steeper borders, and response amplitudes were higher when eye movement compensation was used. In sum, compensating for eye movements facilitated more precise definition of the receptive field. We also monitored horizontal vergence over long sequences of fixation trials and found the variability to be low, as expected for this precise behavior. Our results imply that eye coil signals can be highly accurate and useful for optimizing visual physiology when rigorous precautions are observed.     

Unusually large receptive fields in cats with restricted visual experience

Summary The receptive fields of striate cortex neurons were analyzed in cats which had restricted or no visual experience. Two groups of animals were investigated: 1. cats which were deprived from contour vision over variable periods of time up to 1 year and 2. kittens whose visual experience was restricted to vertically oriented gratings of constant spatial frequency which moved unidirectionally at a fixed distance in front of the restrained animals. In both preparations exceedingly large receptive fields (up to 20° in diameter) were encountered, especially in cells located in supragranular layers. These large receptive fields never extended over more than 2° into the ipsilateral hemifield. Their sensitivity profile was frequently asymmetric and contained discontinuities. Many of these large receptive fields consisted of several excitatory subregions which were separated from each other by as much as 15°. Often but not always the most sensitive area was located where the retinotopic map predicted the receptive field center. The orientation and direction selectivity and also the angular separation of such multiple excitatory bands often matched precisely the orientation, direction and spatial frequency of the experienced moving grating. In other fields with multiple excitatory subregions such a correspondence could not be established; the various subregions could even have different orientation and direction selectivities. From these unconventional receptive fields it is concluded that the function of cat striate cortex is not confined to a point by point analysis of the visual field in retinotopically organized and functionally isolated columns.     

Fast-spiking interneurons have an initial orientation bias that is lost with vision

We found that in mice, following eye opening, fast-spiking, parvalbumin-positive GABAergic interneurons had well-defined orientation tuning preferences and that subsequent visual experience broadened this tuning. Broad inhibitory tuning was not required for the developmental sharpening of excitatory tuning but did precede binocular matching of excitatory orientation tuning. We propose that experience-dependent broadening of inhibition is a candidate for initiating the critical period of excitatory binocular plasticity in developing visual cortex.