Direction Selectivity Index Research Articles

Neuron populations across layer 2-6 in the mouse visual cortex exhibit different coding abilities in the awake mice.

The visual cortex is a key region in the mouse brain, responsible for processing visual information. Comprised of six distinct layers, each with unique neuronal types and connections, the visual cortex exhibits diverse decoding properties across its layers. This study aimed to investigate the relationship between visual stimulus decoding properties and the cortical layers of the visual cortex while considering how this relationship varies across different decoders and brain regions. This study reached the above conclusions by analyzing two publicly available datasets obtained through two-photon microscopy of visual cortex neuronal responses. Various types of decoders were tested for visual cortex decoding. Our findings indicate that the decoding accuracy of neuronal populations with consistent sizes varies among visual cortical layers for visual stimuli such as drift gratings and natural images. In particular, layer 4 neurons in VISp exhibited significantly higher decoding accuracy for visual stimulus identity compared to other layers. However, in VISm, the decoding accuracy of neuronal populations with the same size in layer 2/3 was higher than that in layer 4, despite the overall accuracy being lower than that in VISp and VISl. Furthermore, SVM surpassed other decoders in terms of accuracy, with the variation in decoding performance across layers being consistent among decoders. Additionally, we found that the difference in decoding accuracy across different imaging depths was not associated with the mean orientation selectivity index (OSI) and the mean direction selectivity index (DSI) neurons, but showed a significant positive correlation with the mean reliability and mean signal-to-noise ratio (SNR) of each layer's neuron population. These findings lend new insights into the decoding properties of the visual cortex, highlighting the role of different cortical layers and decoders in determining decoding accuracy. The correlations identified between decoding accuracy and factors such as reliability and SNR pave the way for more nuanced understandings of visual cortex functioning.

Electrically-evoked responses for retinal prostheses are differentially altered depending on ganglion cell types in outer retinal neurodegeneration caused by Crb1 gene mutation.

Microelectronic prostheses for artificial vision stimulate neurons surviving outer retinal neurodegeneration such as retinitis pigmentosa (RP). Yet, the quality of prosthetic vision substantially varies across subjects, maybe due to different levels of retinal degeneration and/or distinct genotypes. Although the RP genotypes are remarkably diverse, prosthetic studies have primarily used retinal degeneration (rd) 1 and 10 mice, which both have Pde6b gene mutation. Here, we report the electric responses arising in retinal ganglion cells (RGCs) of the rd8 mouse model which has Crb1 mutation. We first investigated age-dependent histological changes of wild-type (wt), rd8, and rd10 mice retinas by H&E staining. Then, we used cell-attached patch clamping to record spiking responses of ON, OFF and direction selective (DS) types of RGCs to a 4-ms-long electric pulse. The electric responses of rd8 RGCs were analyzed in comparison with those of wt RGCs in terms of individual RGC spiking patterns, populational characteristics, and spiking consistency across trials. In the histological examination, the rd8 mice showed partial retinal foldings, but the outer nuclear layer thicknesses remained comparable to those of the wt mice, indicating the early-stage of RP. Although spiking patterns of each RGC type seemed similar to those of the wt retinas, correlation levels between electric vs. light response features were different across the two mouse models. For example, in comparisons between light vs. electric response magnitudes, ON/OFF RGCs of the rd8 mice showed the same/opposite correlation polarity with those of wt mice, respectively. Also, the electric response spike counts of DS RGCs in the rd8 retinas showed a positive correlation with their direction selectivity indices (r = 0.40), while those of the wt retinas were negatively correlated (r = -0.90). Lastly, the spiking timing consistencies of late responses were largely decreased in both ON and OFF RGCs in the rd8 than the wt retinas, whereas no significant difference was found across DS RGCs of the two models. Our results indicate the electric response features are altered depending on RGC types even from the early-stage RP caused by Crb1 mutation. Given the various degeneration patterns depending on mutation genes, our study suggests the importance of both genotype- and RGC type-dependent analyses for retinal prosthetic research.

Differences in auditory temporal processing in the left and right auditory cortices of the rat

In the present study, we examined hemispheric differences in the representation and processing of temporally structured auditory stimuli. Neuronal responses evoked by sinusoidally frequency modulated (FM) tones, frequency sweeps, amplitude modulated (AM) tones and noise, click trains with constant inter-click intervals and natural vocalizations were recorded from the left (LAC) and right (RAC) auditory cortices in adult (4–6 months old) anaesthetized F344 rats. Using vector strength, modulation-transfer functions, van Rossum distances, or direction-selectivity index, representation and processing of structured auditory stimuli were compared in the LAC and the RAC. The RAC generally tended to exhibit a higher ability to synchronize with the stimulus, a higher reproducibility of responses, and a higher proportion of direction-selective units. The LAC, on the other hand, mostly had higher relative response magnitudes in the modulation transfer functions. Importantly, the hemispheric differences were dependent on the type of the stimulus and there was also a significant inter-individual variability. Our findings indicate that neural coding in the RAC is based more on timing of action potentials (temporal code), while the LAC uses more the response magnitudes (rate code). It is thus necessary to distinguish between the type of the neural code and the stimulus feature it encodes and reconsider the simple opinion about dominance of the LAC for temporal processing, as it may not hold in general for all types of temporally structured stimuli.

Spatial clustering of orientation preference in primary visual cortex of the large rodent agouti.

SummaryAll rodents investigated so far possess orientation-selective neurons in the primary visual cortex (V1) but – in contrast to carnivores and primates – no evidence of periodic maps with pinwheel-like structures. Theoretical studies debating whether phylogeny or universal principles determine development of pinwheels point to V1 size as a critical constraint. Thus, we set out to study maps of agouti, a big diurnal rodent with a V1 size comparable to cats'. In electrophysiology, we detected interspersed orientation and direction-selective neurons with a bias for horizontal contours, corroborated by homogeneous activation in optical imaging. Compatible with spatial clustering at short distance, nearby neurons tended to exhibit similar orientation preference. Our results argue against V1 size as a key parameter in determining the presence of periodic orientation maps. They are consistent with a phylogenetic influence on the map layout and development, potentially reflecting distinct retinal traits or interspecies differences in cortical circuitry.

A Model for the Origin of Motion Direction Selectivity in Visual Cortex.

Motion perception is a vital part of our sensory repertoire in that it contributes to navigation, awareness of moving objects, and communication. Motion sense in carnivores and primates originates with primary visual cortical neurons selective for motion direction. More than 60 years after the discovery of these neurons, there is still no consensus on the mechanism underlying direction selectivity. This paper describes a model of the cat's visual system in which direction selectivity results from the well-documented orientation selectivity of inhibitory neurons: inhomogeneities in the orientation preference map for inhibitory neurons leads to spatially asymmetric inhibition, and thus to direction selectivity. Stimulation of the model with a drifting grating shows that direction selectivity results from the relative timing of excitatory and inhibitory inputs to a neuron. Using a stationary contrast-reversing grating reveals that the inhibitory input is spatially displaced in the preferred direction relative to the excitatory input, and that this asymmetry leads to the timing difference. More generally, the model yields physiologically realistic estimates of the direction selectivity index, and it reproduces the critical finding with contrast-reversing gratings that response phase advances with grating spatial phase. It is concluded that a model based on intracortical inhibition can account well for the known properties of direction selectivity in carnivores and primates.SIGNIFICANCE STATEMENT Motion perception is vital for navigation, communication, and the awareness of moving objects. Motion sense depends on cortical neurons that are selective for motion direction, and this paper describes a model for the physiological mechanism underlying cortical direction selectivity. The essence of the model is that intracortical inhibition of a direction-selective cell is spatially inhomogeneous and therefore depends on whether a stimulus generates inhibition before or after reaching the cell's receptive field: the response is weaker in the former than in the latter case. If the model is correct, it will contribute to the understanding of motion processing in carnivores and primates.

Silencing "Top-Down" Cortical Signals Affects Spike-Responses of Neurons in Cat's "Intermediate" Visual Cortex.

We examined the effects of reversible inactivation of a higher-order, pattern/form-processing, postero-temporal visual (PTV) cortex on the background activities and spike-responses of single neurons in the ipsilateral cytoarchitectonic area 19 (putative area V3) of anesthetized domestic cats. Very occasionally (2/28), silencing recurrent “feedback” signals from PTV, resulted in significant and reversible reduction in background activity of area 19 neurons. By contrast, in large proportions of area 19 neurons, PTV inactivation resulted in: (i) significant reversible changes in the peak magnitude of their responses to visual stimuli (35.5%; 10/28); (ii) substantial reversible changes in direction selectivity indices (DSIs; 43%; 12/28); and (iii) reversible, upward shifts in preferred stimulus velocities (37%; 7/19). Substantial (≥20°) shifts in preferred orientation and/or substantial (≥20°) changes in width of orientation-tuning curves of area 19 neurons were however less common (26.5%; 4/15). In a series of experiments conducted earlier, inactivation of PTV also induced upward shifts in the preferred velocities of the ipsilateral cytoarchitectonic area 17 (V1) neurons responding optimally at low velocities. These upward shifts in preferred velocities of areas 19 and 17 neurons were often accompanied by substantial increases in DSIs. Thus, in both the primary visual cortex and the “intermediate” visual cortex (area 19), feedback from PTV plays a modulatory role in relation to stimulus velocity preferences and/or direction selectivity, that is, the properties which are usually believed to be determined by the inputs from the dorsal thalamus and/or feedforward inputs from the primary visual cortices. The apparent specialization of area 19 for processing information about stationary/slowly moving visual stimuli is at least partially determined, by the feedback from the higher-order pattern-processing visual area. Overall, the recurrent signals from the higher-order, pattern/form-processing visual cortex appear to play an important role in determining the magnitude of spike-responses and some “motion-related” receptive field properties of a substantial proportion of neurons in the intermediate form-processing visual area—area 19.

Direction selectivity of neurons in the visual cortex is non-linear and lamina-dependent.

Neurons in the visual cortex are generally selective to direction of movement of a stimulus. Although models of this direction selectivity (DS) assume linearity, experimental data show stronger degrees of DS than those predicted by linear models. Our current study was intended to determine the degree of non-linearity of the DS mechanism for cells within different laminae of the cat's primary visual cortex. To do this, we analysed cells in our database by using neurophysiological and histological approaches to quantify non-linear components of DS in four principal cortical laminae (layers 2/3, 4, 5, and 6). We used a DS index (DSI) to quantify degrees of DS in our sample. Our results showed laminar differences. In layer 4, the main thalamic input region, most neurons were of the simple type and showed high DSI values. For complex cells in layer 4, there was a broad distribution of DSI values. Similar features were observed in layer 2/3, but complex cells were dominant. In deeper layers (5 and 6), DSI value distributions were characterized by clear peaks at high values. Independently of specific lamina, high DSI values were accompanied by narrow orientation tuning widths. Differences in orientation tuning for non-preferred vs. preferred directions were smallest inlayer 4 and largest in layer 6. These results are consistent with a non-linear process of intra-cortical inhibition that enhances DS by selective suppression of neuronal firing for non-preferred directions of stimulus motion in a lamina-dependent manner. Other potential mechanisms are also considered.

Weak orientation and direction selectivity in lateral geniculate nucleus representing central vision in the gray squirrel Sciurus carolinensis.

Classic studies of lateral geniculate nucleus (LGN) and visual cortex (V1) in carnivores and primates have found that a majority of neurons in LGN exhibit a center-surround organization, while V1 neurons exhibit strong orientation selectivity and, in many species, direction selectivity. Recent work in the mouse and the monkey has discovered previously unknown classes of orientation- and direction-selective neurons in LGN. Furthermore, some recent studies in the mouse report that many LGN cells exhibit pronounced orientation biases that are of comparable strength to the subthreshold inputs to V1 neurons. These results raise the possibility that, in rodents, orientation biases of individual LGN cells make a substantial contribution to cortical orientation selectivity. Alternatively, the size and contribution of orientation- or direction-selective channels from LGN to V1 may vary across mammals. To address this question, we examined orientation and direction selectivity in LGN and V1 neurons of a highly visual diurnal rodent: the gray squirrel. In the representation of central vision, only a few LGN neurons exhibited strong orientation or direction selectivity. Across the population, LGN neurons showed weak orientation biases and were much less selective for orientation compared with V1 neurons. Although direction selectivity was weak overall, LGN layers 3abc, which contain neurons that express calbindin, exhibited elevated direction selectivity index values compared with LGN layers 1 and 2. These results suggest that, for central visual fields, the contribution of orientation- and direction-selective channels from the LGN to V1 is small in the squirrel. As in other mammals, this small contribution is elevated in the calbindin-positive layers of the LGN.

Neurons in cat V1 show significant clustering by degree of tuning.

Neighboring neurons in cat primary visual cortex (V1) have similar preferred orientation, direction, and spatial frequency. How diverse is their degree of tuning for these properties? To address this, we used single-tetrode recordings to simultaneously isolate multiple cells at single recording sites and record their responses to flashed and drifting gratings of multiple orientations, spatial frequencies, and, for drifting gratings, directions. Orientation tuning width, spatial frequency tuning width, and direction selectivity index (DSI) all showed significant clustering: pairs of neurons recorded at a single site were significantly more similar in each of these properties than pairs of neurons from different recording sites. The strength of the clustering was generally modest. The percent decrease in the median difference between pairs from the same site, relative to pairs from different sites, was as follows: for different measures of orientation tuning width, 29-35% (drifting gratings) or 15-25% (flashed gratings); for DSI, 24%; and for spatial frequency tuning width measured in octaves, 8% (drifting gratings). The clusterings of all of these measures were much weaker than for preferred orientation (68% decrease) but comparable to that seen for preferred spatial frequency in response to drifting gratings (26%). For the above properties, little difference in clustering was seen between simple and complex cells. In studies of spatial frequency tuning to flashed gratings, strong clustering was seen among simple-cell pairs for tuning width (70% decrease) and preferred frequency (71% decrease), whereas no clustering was seen for simple-complex or complex-complex cell pairs.

Binocular neurons in parastriate cortex: interocular 'matching' of receptive field properties, eye dominance and strength of silent suppression.

Spike-responses of single binocular neurons were recorded from a distinct part of primary visual cortex, the parastriate cortex (cytoarchitectonic area 18) of anaesthetized and immobilized domestic cats. Functional identification of neurons was based on the ratios of phase-variant (F1) component to the mean firing rate (F0) of their spike-responses to optimized (orientation, direction, spatial and temporal frequencies and size) sine-wave-luminance-modulated drifting grating patches presented separately via each eye. In over 95% of neurons, the interocular differences in the phase-sensitivities (differences in F1/F0 spike-response ratios) were small (≤0.3) and in over 80% of neurons, the interocular differences in preferred orientations were ≤10°. The interocular correlations of the direction selectivity indices and optimal spatial frequencies, like those of the phase sensitivies and optimal orientations, were also strong (coefficients of correlation r ≥0.7005). By contrast, the interocular correlations of the optimal temporal frequencies, the diameters of summation areas of the excitatory responses and suppression indices were weak (coefficients of correlation r ≤0.4585). In cells with high eye dominance indices (HEDI cells), the mean magnitudes of suppressions evoked by stimulation of silent, extra-classical receptive fields via the non-dominant eyes, were significantly greater than those when the stimuli were presented via the dominant eyes. We argue that the well documented ‘eye-origin specific’ segregation of the lateral geniculate inputs underpinning distinct eye dominance columns in primary visual cortices of mammals with frontally positioned eyes (distinct eye dominance columns), combined with significant interocular differences in the strength of silent suppressive fields, putatively contribute to binocular stereoscopic vision.

Anterior-Posterior Direction Opponency in the Superficial Mouse Lateral Geniculate Nucleus

We show functional-anatomical organization of motion direction in mouse dorsal lateral geniculate nucleus (dLGN) using two-photon calcium imaging of dense populations in thalamus. Surprisingly, thesuperficial 75μm region contains anterior and posterior direction-selective neurons (DSLGNs) intermingled with nondirection-selective neurons, while upward- and downward-selective neurons are nearlyabsent. Unexpectedly, the remaining neurons encode both anterior and posterior directions, forming horizontal motion-axis selectivity. A model of random wiring consistent with these results makes quantitative predictions about the connectivity of direction-selective retinal ganglion cell (DSRGC) inputs to the superficial dLGN. DSLGNs are more sharply tuned than DSRGCs. These results suggest that dLGN maintains and sharpens retinal direction selectivity and integrates opposing DSRGC subtypes in a functional-anatomical region, perhaps forming a feature representation for horizontal-axis motion, contrary to dLGN being a simple relay. Furthermore, they support recent conjecture that cortical direction and orientation selectivity emerge in part from a previously undescribed motion-selective retinogeniculate pathway.

Visual Pathway for the Optokinetic Reflex in Infant Macaque Monkeys

The horizontal optokinetic nystagmus (hOKN) in primates is immature at birth. To elucidate the early functional state of the visual pathway for hOKN, retinal slip neurons were recorded in the nucleus of the optic tract and dorsal terminal nucleus (NOT-DTN) of 4 anesthetized infant macaques. These neurons were direction selective for ipsiversive stimulus movement shortly after birth [postnatal day 9 (P9)], although at a lower direction selectivity index (DSI). The DSI in the older infants (P12, P14, P60) was not different from adults. A total of 96% of NOT-DTN neurons in P9, P12, and P14 were binocular, however, significantly more often dominated by the contralateral eye than in adults. Already in the youngest animals, NOT-DTN neurons were well tuned to different stimulus velocities; however, tuning was truncated toward lower stimulus velocities when compared with adults. As early as at P12, electrical stimulation in V1 elicited orthodromic responses in the NOT-DTN. However, the incidence of activated neurons was much lower in infants (40-60% of the tested NOT-DTN neurons) than in adults (97%). Orthodromic latencies from V1 were significantly longer in P12-P14 (x = 12.2 ± 8.9 ms) than in adults (x = 3.51 ± 0.81 ms). At the same age, electrical stimulation in motion-sensitive area MT was more efficient in activating NOT-DTN neurons (80% of the tested cells) and yielded shorter latencies than in V1 (x = 7.8 ± 3.02 ms; adult x = 2.99 ± 0.85 ms). The differences in discharge rate between neurons in the NOT-DTN contra- and ipsilateral to the stimulated eye are equivalent to the gain asymmetry between monocularly elicited OKN in temporonasal and nasotemporal direction at the various ages.

Region-specificity of GABAA receptor mediated effects on orientation and direction selectivity in cat visual cortical area 18

The role of GABAergic inhibition in orientation and direction selectivity has been investigated with the GABA(A)-Blocker bicuculline in the cat visual cortex, and results indicated a region specific difference of functional contributions of GABAergic inhibition in areas 17 and 18. In area 17 inhibition appeared mainly involved in sculpturing orientation and direction tuning, while in area 18 inhibition seemed more closely associated with temporal receptive field properties. However, different types of stimuli were used to test areas 17 and 18 and further studies performed in area 17 suggested an important influence of the stimulus type (single light bars vs. moving gratings) on the evoked responses (transient vs. sustained) and inhibitory mechanisms (GABA(A) vs. GABA(B)) which in turn might be more decisive for the specific results than the cortical region. To insert the missing link in this chain of arguments it was necessary to study GABAergic inhibition in area 18 with moving light bars, which has not been done so far. Therefore, in the present study we investigated area 18 cells responding to oriented moving light bars with extracellular recordings and reversible microiontophoretic blockade of GABAergig inhibition with bicuculline methiodide. The majority of neurons was characterized by a pronounced orientation specificity and variable degrees of direction selectivity. GABA(A)ergic inhibition significantly influenced preferred orientation and preferred direction in area 18. During the action of bicuculline orientation tuning width increased and orientation and direction selectivity indices decreased. Our results obtained in area 18 with moving bar stimuli, although in the proportion of affected cells similar to those described in area 17, quantitatively matched the findings for direction and orientation specificity obtained with moving gratings in area 18. Accordingly, stimulus type is not decisive in area 18 and the GABA(A) dependent, inhibitory intracortical computations involved in orientation specificity are indeed region-specific and in comparison to area 17 less effective in area 18.

Motor Cortex Gates Vibrissal Responses in a Thalamocortical Projection Pathway

Higher-order thalamic nuclei receive input from both the cerebral cortex and prethalamic sensory pathways. However, at rest these nuclei appear silent due to inhibitory input from extrathalamic regions, and it has therefore remained unclear how sensory gating of these nuclei takes place. In the rodent, the ventral division of the zona incerta (ZIv) serves as a relay station within the paralemniscal thalamocortical projection pathway for whisker-driven motor activity. Most, perhaps all, ZIv neurons are GABAergic, and recent studies have shown that these cells participate in a feedforward inhibitory circuit that blocks sensory transmission in the thalamus. The present study provides evidence that the stimulation of the vibrissa motor cortex suppresses vibrissal responses in ZIv via an intra-incertal GABAergic circuit. These results provide support for the proposal that sensory transmission operates via a top-down disinhibitory mechanism that is contingent on motor activity.

The effects of reversible inactivation of postero-temporal visual cortex on neuronal activities in cat's area 17

‘Spontaneous’ and visually evoked action potentials were recorded from single neurons in cytoarchitectonic area 17 (striate cortex, area V1) of anaesthetized and immobilized cats, prior to, during and after brief reversible inactivation of the ipsilateral postero-temporal visual (PTV) cortex (presumed homologue of primate inferotemporal cortex). Inactivation of PTV cortex resulted: 1) in significant changes in the response magnitude (mostly a reduction) to optimal and/or sub-optimal visual stimuli in over 55% of area 17 cells and 2) significant changes (usually a reduction) in the ‘spontaneous’ (background) activity of about two-thirds of the cells in which inactivation of PTV cortex significantly affected the magnitude of responses to optimal stimuli. In over 85% of the significantly affected area 17 cells, rewarming PTV cortex to normal temperature (36 °C) resulted in the recovery of both the magnitude of responses and the background activity to levels not significantly different from pre-inactivation levels. Irrespective of the significance of changes in the magnitude of responses, in a substantial proportion of area 17 cells, inactivation of PTV cortex resulted in changes in some receptive field characteristics. Thus, there were substantial (20% or more) changes in orientation tuning widths (in over a quarter of the sample) and/or direction selectivity indices (in about a third of the sample). Thus, the feedback signals originating from PTV cortex, like signals originating from some other ‘higher-order’ visual cortical areas exert a clear modulatory influence on the responsiveness, background activity and some receptive field properties of neurons in the ipsilateral area 17.

Area 21a of cat visual cortex strongly modulates neuronal activities in the superior colliculus.

We have examined the influence of cortico-tectal projections from one of the pattern-processing extrastriate visual cortical areas, area 21a, on the responses to visual stimuli of single neurones in the superior colliculi of adult cats. For this purpose area 21a was briefly inactivated by cooling to 10 degrees C using a Peltier device. Responses to visual stimuli before and during cooling as well as after rewarming ipsilateral area 21a were compared. In addition, in a subpopulation of collicular neurones we have studied the effects of reversible inactivation of ipsilateral striate cortex (area 17, area V1). When area 21a was cooled, the temperature of area 17 was kept at 36 degrees C and vice versa. In the majority of cases (41/65; 63%), irrespective of the velocity response profiles of collicular neurones, inactivation of area 21a resulted in a significant decrease in magnitude of responses of neurones in the ipsilateral colliculus and only in a small proportion of cells (2/65; 3.1%) was there a significant increase in the magnitude of responses. Inactivation of area 21a resulted in significant changes in the magnitude of responses of collicular cells located not only in the retino-recipient layers but also in the stratum griseum intermediale. In most cases, reversible inactivation of area 17 resulted in a greater reduction in the magnitude of responses of collicular cells than inactivation of area 21a. Reversible inactivation of area 21a also affected the direction selectivity indices and length tuning of most collicular cells tested.

The spatial substructure of visual receptive fields in the cat's superior colliculus.

Although the direction selective properties of the superficial layer cells of the cat's superior colliculus have been extensively studied, the mechanisms underlying this property remain controversial. With the aim to understand the mechanism(s) underlying directional selectivity of collicular neurons we examined the substructure of their visual receptive fields. 1. The strength of cell responses and the direction selectivity indices varied in relation to the location of the tested region within the receptive field and the amplitude of stimulus movement. 2. Decrease of the amplitude of motion resulted in a decrease of direction selectivity index both in the group of direction-selective cells and in the group of cells classified as direction nonselective but with a directional bias. 3. The decrease of direction selectivity for small amplitude movement resulted mainly from increase in the magnitude of response in the nonpreferred direction of movement. 4. These results suggest that the receptive fields of most collicular cells are composed of subregions with different response profiles and indicate that inhibitory mechanisms dictate direction selectivity of collicular cells.

Excitatory convergence of Y and non-Y channels onto single neurons in the anterior ectosylvian visual area of the cat.

Numerous functional and hodological studies of the anterior ectosylvian visual area (AEV) of the cerebral cortex of the cat suggest that this area plays an important role in processing information about visual motion. In the present study, in cats with selective conduction block of Y fibres in one optic nerve, we have examined the extent of the excitatory convergence of Y (presumed 'motion channel') and non-Y information channels on single neurons in AEV, as well as the contribution of the Y channel to the receptive field properties of AEV neurons. While in normal cats all neurons recorded from AEV were binocular, i.e. could be photically activated via either eye, in cats with selective conduction block of Y fibres in one optic nerve, a significant proportion (about 15%) of AEV cells could be photically activated only via the normal eye. In comparison to those in normal cats, the peak discharge rates of AEV neurons in the Y-blocked cats were drastically reduced not only when photic stimuli were presented via the Y-blocked eye, but also when they were presented via the normal eye. Selective block of Y input also resulted in a significant shift in velocity preferences towards the lower velocities. However, the direction selectivity indices of AEV neurons were not affected by selective Y block. Thus: (i) the responses of AEV neurons to a high velocity of motion are dependent on the integrity of the Y input; (ii) the 'spontaneous' (i.e. not photically evoked) discharges of Y retinal ganglion cells exert a facilitatory influence on the responses of AEV cells to photic stimuli; (iii) although the responses of AEV neurons are dominated by the Y inputs, AEV neurons also receive significant non-Y excitatory inputs; and (iv) the strong direction selectivity revealed in most AEV neurons does not dependent on the integrity of Y input.

Excitatory convergence of Y and non-Y information channels on single neurons in the PMLS area, a motion area of the cat visual cortex.

We analysed the receptive field properties of neurons in the posteromedial lateral suprasylvian (PMLS) visual cortical area of anaesthetized cats in which there was selective conduction block of the largest (Y-type) fibres in one optic nerve. As in normal cats, in cats with selective block of one optic nerve the great majority of PMLS cells could be activated by photic stimulation through either eye. However, the responses evoked by stimulation via the eye with the selectively pressure-blocked optic nerve ('Y-blocked eye') were significantly weaker than those of the same cells evoked by the stimulation via the normal eye. Accordingly, eye dominance histograms were shifted markedly in favour of the normal eye. Furthermore, there was a significant shift towards lower preferred velocities when PMLS cells were photically stimulated via the Y-blocked eye. Finally, when stimulated via the Y-blocked eye, PMLS cells responded poorly or not at all to high stimulus velocities (> or = 100 degrees/s). On the other hand, a number of receptive field properties, such as the spatial organization of receptive fields, the size of the discharge fields, orientation tuning and direction selectivity indices, were not significantly affected by the removal of the Y input. We conclude that virtually all neurons in the PMLS area of the cat receive excitatory input from both Y and non-Y information channels, although the Y channel provides the dominant input and makes the principal contribution to the detection of high-velocity motion.

Areas PMLS and 21a of cat visual cortex: two functionally distinct areas.

We have compared the receptive field properties of neurons recorded from visuotopically corresponding regions of area 21a and the posteromedial lateral suprasylvian area (PMLS) of cat visual cortex. In both areas, the great majority of neurons were orientation-selective and binocular, and their responses to moving contours were modulated by simultaneous in-phase or anti-phase motion of large textured background stimuli ('visual noise'). However, despite the great hodological similarity between the two areas, PMLS neurons had on average significantly higher peak discharge rates, exhibited substantially greater direction selectivity indices, and preferred substantially higher stimulus velocities than area 21a neurons. Furthermore, the majority of binocular neurons in the PMLS area and in area 21a were dominated respectively by contralateral and ipsilateral eyes. Finally, while 46% of PMLS neurons were excited by movement of visual noise per se, only 25% of area 21a neurons could be excited by such stimuli. We argue that the PMLS area, like its presumed primate homologue the middle-temporal (MT) area, is mainly involved in motion analysis. By contrast, area 21a appears to be involved in pattern analysis rather than motion analysis. It is likely that phylogenetically area 21a derives from the PMLS area.