Comparison of recordings from microelectrode arrays and single electrodes in the visual cortex.
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Action Potentials↗
Biomedical subjects
A B Bonds
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Synchronous activity in cat visual cortex encodes collinear and cocircular contours.
We explored how contour information in primary visual cortex might be embedded in the simultaneous activity of multiple cells recorded with a 100-electrode array. Synchronous activity in cat visual cortex was more selective and predictable in discriminating between drifting grating and concentric ring stimuli than changes in firing rate. Synchrony was found even between cells with wholly different orientation preferences when their receptive fields were circularly aligned, and membership in synchronous groups was orientation and curvature dependent. The existence of synchrony between cocircular cells reinforces its role as a general mechanism for contour integration and shape detection as predicted by association field concepts. Our data suggest that cortical synchrony results from common and synchronous input from earlier visual areas and that it could serve to shape extrastriate response selectivity.
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Real-time visualization of neural synchrony for identifying coordinated cell assemblies.
We introduce a synchrony map that translates the fine temporal organization of multi-unit responses in the visual cortex into an easily interpreted spatial display. We test the synchrony map on microelectrode array recordings in Area 17 of anesthetized and paralyzed cats. We first examine the synchrony map using averaged data and probability calculations to demonstrate orientation-dependent changes in synchrony. We then demonstrate how the synchrony map can be implemented for real-time visualization of synchrony among neural assemblies.
Action Potentials↗
Gamma oscillation maintains stimulus structure-dependent synchronization in cat visual cortex.
Visual cortical cells demonstrate both oscillation and synchronization, although the underlying causes and functional significance of these behaviors remain uncertain. We simultaneously recorded single-unit activity with microelectrode arrays in supragranular layers of area 17 of cats paralyzed and anesthetized with propofol and N(2)O. Rate-normalized autocorrelograms of 24 cells reveal bursting (100%) and gamma oscillation (63%). Renewal density analysis, used to explore the source of oscillation, suggests a contribution from extrinsic influences such as feedback. However, a bursting refractory period, presumably membrane-based, could also encourage oscillatory firing. When we investigated the source of synchronization for 60 cell pairs we found only moderate correlation of synchrony with bursts and oscillation. We did, nonetheless, discover a possible functional role for oscillation. In all cases of cross-correlograms that exhibited oscillation, the strength of the synchrony was maintained throughout the stimulation period. When no oscillation was apparent, 75% of the cell pairs showed decay in their synchronization. The synchrony between cells is strongly dependent on similar response onset latencies. We therefore propose that structured input, which yields tight organization of latency, is a more likely candidate for the source of synchronization than oscillation. The reliable synchrony at response onset could be driven by spatial and temporal correlation of the stimulus that is preserved through the earlier stages of the visual system. Oscillation then contributes to maintenance of the synchrony to enhance reliable transmission of the information for higher cognitive processing.
Action Potentials↗
Cooperative synchronized assemblies enhance orientation discrimination.
There is no clear link between the broad tuning of single neurons and the fine behavioral capabilities of orientation discrimination. We recorded from populations of cells in the cat visual cortex (area 17) to examine whether the joint activity of cells can support finer discrimination than found in individual responses. Analysis of joint firing yields a substantial advantage (i.e., cooperation) in fine-angle discrimination. This cooperation increases to more considerable levels as the population of an assembly is increased. The cooperation in a population of six cells provides encoding of orientation with an information advantage that is at least 2-fold in terms of requiring either fewer cells or less time than independent coding. This cooperation suggests that correlated or synchronized activity can increase information.
Animals↗
From another angle: Differences in cortical coding between fine and coarse discrimination of orientation.
We measured the information available for orientation discrimination from metric distances for 24 cells in area 17 of cats that were paralyzed and anesthetized with Propofol and N(2)O. The metric distance information confirms fundamental coding differences for discrimination between fine (<10 degrees ) and coarse (>10 degrees ) orientation differences. The information for discriminating larger orientation differences is contained mainly in the firing rate, with minor enhancements from the coarse (30-70 ms) temporal structure in the firing rate. Both precise spike timing (9.2 ms) and intervals (6.8 ms) sustained over the stimulus presentation provide information for fine discrimination of orientation, where almost no reliable information is provided by the spike count. We compare and confirm the results (using the same data set) to vector distances based on classification theory. The results support a dynamic spiking mechanism where coordinated activity could provide fast and reliable information about detailed angle and/or direction information in the region of the preferred orientation.
Animals↗
Cooperation between area 17 neuron pairs enhances fine discrimination of orientation.
We examined 66 complex cells in area 17 of cats that were paralyzed and anesthetized with propofol and N2O. We studied changes in ensemble responses for small (<10 degrees ) and large (>10 degrees ) differences in orientation. Examination of temporal resolution and discharge history revealed advantages in discrimination from both dependent (e.g., synchronization) and independent (e.g., bursting) interspike interval properties. For 27 pairs of neurons, we found that the average cooperation (the advantage gained from the joint activity) was 57.6% for fine discrimination of orientation but <5% for gross discrimination. Dependency (probabilistic quantification of the interaction between the cells) was measured between 29 pairs of neurons while varying orientation. On average, the dependency tuning for orientation was 35.5% narrower than the average firing rate tuning. The changes in dependency around the peak orientation (at which the firing rate remains relatively constant) lead to substantial cooperation that can improve discrimination in this region. The narrow tuning of dependency and the cooperation provide evidence to support a population-encoding scheme that is based on biologically plausible mechanisms and that could account for hyperacuities.
Action Potentials↗
A comparison of koniocellular, magnocellular and parvocellular receptive field properties in the lateral geniculate nucleus of the owl monkey (Aotus trivirgatus).
1. By analogy to previous work on lateral geniculate nucleus (LGN) magnocellular (M) and parvocellular (P) cells our goal was to construct a physiological profile of koniocellular (K) cells that might be linked to particular visual perceptual attributes. 2. Extracellular recordings were used to study LGN cells, or their axons, in silenced primary visual cortex (V1), in nine anaesthetized owl monkeys injected with a neuromuscular blocker. Receptive field centre-surround organization was examined using flashing spots. Spatial and temporal tuning and contrast responses were examined using drifting sine-wave gratings; counterphase sine-wave gratings were used to examine linearity of spatial summation. 3. Receptive fields of 133 LGN cells and 10 LGN afferent axons were analysed at eccentricities ranging from 2.8 to 31.3 deg. Thirty-four per cent of K cells and only 9 % of P and 6 % of M cells responded poorly to drifting gratings. K, P and M cells showed increases in centre size with eccentricity, but K cells showed more scatter. All cells, except one M cell, showed linearity in spatial summation. 4. At matched eccentricities, K cells exhibited lower spatial and intermediate temporal resolution compared with P and M cells. K contrast thresholds and gains were more similar to those of M than P cells. M cells showed lower spatial and higher temporal resolution and contrast gains than P cells. 5. K cells in different K LGN layers differed in spatial, temporal and contrast characteristics, with K3 cells having higher spatial resolution and lower temporal resolution than K1/K2 cells. 6. Taken together with previous results these findings suggest that the K cells consist of several classes, some of which could contribute to conventional aspects of spatial and temporal resolution.
Algorithms↗
Modification of response functions of cat visual cortical cells by spatially congruent perturbing stimuli.
Responses of cat striate cortical cells to a drifting sinusoidal grating were modified by the superimposition of a second, perturbing grating (PG) that did not excite the cell when presented alone. One consequence of the presence of a PG was a shift in the tuning curves. The orientation tuning of all 41 cells exposed to a PG and the spatial frequency tuning of 83% of the 23 cells exposed to a PG showed statistically significant dislocations of both the response function peak and center of mass from their single grating values. As found in earlier reports, the presence of PGs suppressed responsiveness. However, reductions measured at the single grating optimum orientation or spatial frequency were on average 1.3 times greater than the suppression found at the peak of the response function modified by the presence of the PG. Much of the loss in response seen at the single grating optimum is thus a result of a shift in the tuning function rather than outright suppression. On average orientation shifts were repulsive and proportional (approximately 0.10 deg/deg) to the angle between the perturbing stimulus and the optimum single grating orientation. Shifts in the spatial frequency response function were both attractive and repulsive, resulting in an overall average of zero. For both simple and complex cells, PGs generally broadened orientation response function bandwidths. Similarly, complex cell spatial frequency response function bandwidths broadened. Simple cell spatial frequency response functions usually did not change, and those that did broadened only 4% on average. These data support the hypothesis that additional sinusoidal components in compound stimuli retune cells' response functions for orientation and spatial frequency.
Algorithms↗
Classification of non-stationary neural signals.
Although a number of methods have been proposed for classification of individual action potentials embedded in multi-unit activity, they have been challenged by non-stationarity. The waveform shapes of action potentials can change rapidly over time as a result of shifts in membrane conductances during extended burst firing sequences and more slowly over time due to electrode drift. These changes are typically non-Gaussian. We present an algorithm for waveform identification that makes no assumptions on the distribution of these shapes other than the change in waveform shape for a particular neuron should not be discontinuous. We apply this algorithm to the resolution of multi-unit neural signals recorded in the cat visual cortex and we compare this approach to a spike sorting method that is based on the Bayesian likelihood of a spike fitting a particular model (Lewicki, M. Bayesian modeling and classification of neural signals. Neural Comput 1994;6(5):1005-1030.
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Burst firing and modulation of functional connectivity in cat striate cortex.
We studied the influences of the temporal firing patterns of presynaptic cat visual cortical cells on spike generation by postsynaptic cells. Multiunit recordings were dissected into the activity of individual neurons within the recorded group. Cross-correlation analysis was then used to identify directly coupled neuron pairs. The 22 multiunit groups recorded typically showed activity from two to six neurons, each containing between 1 and 15 neuron pairs. From a total of 241 neuron pairs, 91 (38%) had a shifted cross-correlation peak, which indicated a possible direct connection. Only two multiunit groups contained no shifted peaks. Burst activity, defined by groups of two or more spikes with intervals of </=8 ms from any single neuron, was analyzed in terms of its effectiveness in eliciting a spike from a second, driven neuron. We defined effectiveness as the percentage of spikes from the driving neuron that are time related to spikes of the driven neuron. The effectiveness of bursts (of any length) in eliciting a time-related response spike averaged 18.53% across all measurements as compared with the effectiveness of single spikes, which averaged 9.53%. Longer bursts were more effective than shorter ones. Effectiveness was reduced with spatially nonoptimal, as opposed to optimal, stimuli. The effectiveness of both bursts and single spikes decreased by the same amount across measurements with nonoptimal orientations, spatial frequencies and contrasts. At similar firing rates and burst lengths, the decrease was more pronounced for nonoptimal orientations than for lower contrasts, suggesting the existence of a mechanism that reduces effectiveness at nonoptimal orientations. These results support the hypothesis that neural information can be emphasized via instantaneous rate coding that is not preserved over long intervals or over trials. This is consistent with the integrate and fire model, where bursts participate in temporal integration.
Action Potentials↗
Stimulus-dependent modulation of spike burst length in cat striate cortical cells.
Burst activity, defined by groups of two or more spikes with intervals of < or = 8 ms, was analyzed in responses to drifting sinewave gratings elicited from striate cortical neurons in anesthetized cats. Bursting varied broadly across a population of 507 simple and complex cells. Half of this population had > or = 42% of their spikes contained in bursts. The fraction of spikes in bursts did not vary as a function of average firing rate and was stationary over time. Peaks in the interspike interval histograms were found at both 3-5 ms and 10-30 ms. In many cells the locations of these peaks were independent of firing rate, indicating a quantized control of firing behavior at two different time scales. The activity at the shorter time scale most likely results from intrinsic properties of the cell membrane, and that at the longer scale from recurrent network excitation. Burst frequency (bursts per s) and burst length (spikes per burst) both depended on firing rate. Burst frequency was essentially linear with firing rate, whereas burst length was a nonlinear function of firing rate and was also governed by stimulus orientation. At a given firing rate, burst length was greater for optimal orientations than for nonoptimal orientations. No organized orientation dependence was seen in bursts from lateral geniculate nucleus cells. Activation of cortical contrast gain control at low response amplitudes resulted in no burst length modulation, but burst shortening at optimal orientations was found in responses characterized by supersaturation. At a given firing rate, cortical burst length was shortened by microinjection of gamma-aminobutyric acid (GABA), and bursts became longer in the presence of N-methyl-bicuculline, a GABA(A) receptor blocker. These results are consistent with a model in which responses are reduced at nonoptimal orientations, at least in part, by burst shortening that is mediated by GABA. A similar mechanism contributes to response supersaturation at high contrasts via recruitment of inhibitory responses that are tuned to adjacent orientations. Burst length modulation can serve as a form of coding by supporting dynamic, stimulus-dependent reorganization of the effectiveness of individual network connections.
Action Potentials↗
Dynamic differentiation of GABAA-sensitive influences on orientation selectivity of complex cells in the cat striate cortex.
The influence of GABAA receptors on orientation selectivity of cat complex cells was tested by iontophoresis of the GABAA receptor blockers bicuculline and N-methyl-bicuculline while stimulating with drifting sinusoidal gratings. Reduction of orientation tuning was markedly less than reported in previous studies that used drifting bars as visual stimuli. Only 3/31 cells lost orientation selectivity, with an average increase in bandwidth of 33%, as opposed to half the cells losing selectivity and a bandwidth increase for the remainder of 47% as reported previously. Infusion of GABAA blockers revealed a prominent stimulus onset transient response, lasting about 120 ms, that showed a broadening of orientation selectivity comparable to that found using drifting bars under similar circumstances. We believe that drifting gratings emphasize a steady-state response component that retains, in the presence of GABAA blockers, significant orientation selectivity. Because the onset transient is initially unselective for orientation, we suggest that the steady-state, orientation-selective response component develops from an alternate inhibitory mechanism, possibly mediated by GABAB receptors.
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Inactivation of the infragranular striate cortex broadens orientation tuning of supragranular visual neurons in the cat.
Intracortical inhibition is believed to enhance the orientation tuning of striate cortical neurons, but the origin of this inhibition is unclear. To examine the possible influence of ascending inhibitory projections from the infragranular layers of striate cortex on the orientation selectivity of neurons in the supragranular layers, we measured the spatiotemporal response properties of 32 supragranular neurons in the cat before, during, and after neural activity in the infragranular layers beneath the recorded cells was inactivated by iontophoretic administration of GABA. During GABA iontophoresis, the orientation tuning bandwidth of 15 (46.9%) supragranular neurons broadened as a result of increases in response amplitude to stimuli oriented about +/- 20 degrees away from the preferred stimulus angle. The mean (+/- SD) baseline orientation tuning bandwidth (half width at half height) of these neurons was 13.08 +/- 2.3 degrees. Their mean tuning bandwidth during inactivation of the infragranular layers increased to 19.59 +/- 2.54 degrees, an increase of 49.7%. The mean percentage increase in orientation tuning bandwidth of the individual neurons was 47.4%. Four neurons exhibited symmetrical changes in their orientation tuning functions, while 11 neurons displayed asymmetrical changes. The change in form of the orientation tuning functions appeared to depend on the relative vertical alignment of the recorded neuron and the infragranular region of inactivation. Neurons located in close vertical register with the inactivated infragranular tissue exhibited symmetric changes in their orientation tuning functions. The neurons exhibiting asymmetric changes in their orientation tuning functions were located just outside the vertical register. Eight of these 11 neurons also demonstrated a mean shift of 6.67 +/- 5.77 degrees in their preferred stimulus orientation. The magnitude of change in the orientation tuning functions increased as the delivery of GABA was prolonged. Responses returned to normal approximately 30 min after the delivery of GABA was discontinued. We conclude that inhibitory projections from neurons within the infragranular layers of striate cortex in cats can enhance the orientation selectivity of supragranular striate cortical neurons.
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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)
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Depression of the cat cortical visual evoked potential by soman.
The effects of intravenous administration of the anticholinesterase agent soman (pinacolyl methylphosphonofluoridate, 3-15 micrograms/kg) on the visual evoked potential (VEP) were examined in cats using phase-reversed sine wave grating stimuli of different spatial frequencies and contrasts. Doses of 5-7 micrograms/kg caused a depression of the VEP across all spatial frequencies in an abrupt, non-graded fashion. Studies in which contrast was varied showed that VEP depression resulted primarily from a decrease in the system gain rather than a change in the contrast sensitivity, and that response depression increased with increasing contrast. The dominant changes in gain revealed by these studies are consistent with a modulation of potassium conductance in the cell membrane which previous studies have shown to be dependent on a cholinergic mechanism.
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Pretreatment with physostigmine, mecamylamine and atropine reduces the impact of soman on the cortical visual evoked potential of the cat.
A pretreatment regimen of physostigmine, mecamylamine and atropine was evaluated for its ability to alleviate the impact of soman on visual system function as measured by changes in the cortical visual evoked potential (VEP) of the cat. Data from unprotected animals showed a threshold (30% depression in the VEP) of 6.4 micrograms/kg, while in pretreated animals, the threshold dose was 32.7 micrograms/kg, yielding a protection ratio of 5:1. Extending the time between pretreatment and exposure reduced the degree of protection. Pretreatment also reduced the degree of VEP depression at suprathreshold doses, indicating a therapeutic effect even in cases of severe exposure.
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Classifying simple and complex cells on the basis of response modulation.
Hubel and Wiesel (1962; Journal of Physiology, London, 160, 106-154) introduced the classification of cortical neurons as simple and complex on the basis of four tests of their receptive field structure. These tests are partly subjective and no one of them unequivocally places neurons into distinct classes. A simple, objective classification criterion based on the form of the response to drifting sinusoidal gratings has been used by several laboratories, although it has been criticized by others. We review published and unpublished evidence which indicates that this simple and objective criterion reliability divides neurons of the striate cortex in both cats and monkeys into two groups that correspond closely to the classically-described simple and complex classes.
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