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D J Tolhurst

Publications and source records attributed to D J Tolhurst.

At least 55 records · Page 3Linked to original sources

The dependence of response amplitude and variance of cat visual cortical neurones on stimulus contrast.

For neurones in the cat's striate cortex, we examined the dependence of response on the contrast of moving sinusoidal gratings. Most neurones showed a clear threshold contrast below which no response was elicited. Such thresholds presumably contribute to the animal's behavioural threshold, which should not be accounted for solely in terms of the detection of a signal in the presence of spontaneous "noise". Above threshold, the response amplitude usually increased linearly with contrast until it began to saturate at the highest contrasts. The variance of the response increased with its amplitude; this finding perhaps underlies the Weber-Fechner relation for psychophysical contrast discrimination.

Animals↗

Non-linearities of temporal summation in neurones in area 17 of the cat.

Sinusoidal temporal modulation of the contrast of sinusoidal gratings has been used to provide one description of the temporal properties of neurones in area 17 of the cat's visual cortex. Concurrently, the waveforms of the neurones' responses to 2 s flashes of identical gratings were examined. In most neurones, the response to the long flash was much more transient than would be expected from the shape of the temporal-frequency tuning curve. Temporal summation in cortical neurones is non-linear.

Animals↗

Spatial summation in the receptive fields of simple cells in the cat's striate cortex.

1. We have examined the responses of simple cells in the cat's atriate cortex to visual patterns that were designed to reveal the extent to which these cells may be considered to sum light-evoked influences linearly across their receptive fields. We used one-dimensional luminance-modulated bars and grating as stimuli; their orientation was always the same as the preferred orientation of the neurone under study. The stimuli were presented on an oscilloscope screen by a digital computer, which also accumulated neuronal responses and controlled a randomized sequence of stimulus presentations. 2. The majority of simple cells respond to sinusoidal gratings that are moving or whose contrast is modulated in time in a manner consistent with the hypothesis that they have linear spatial summation. Their responses to moving gratings of all spatial frequencies are modulated in synchrony with the passage of the gratings' bars across their receptive fields, and they do not produce unmodulated responses even at the highest spatial frequencies. Many of these cells respond to temporally modulated stationary gratings simply by changing their response amplitude sinusoidally as the spatial phase of the grating the grating is varied. Nonetheless, their behavior appears to indicate linear spatial summation, since we show in an Appendix that the absence of a 'null' phase in a visual neurone need not indicate non-linear spatial summation, and further that a linear neurone lacking a 'null' phase should give responses of the form that we have observed in this type of simple cell. 3. A minority of simple cells appears to have significant non-linearities of spatial summation. These neurones respond to moving gratings of high spatial frequency with a partially or totally unmodulated elevation of firing rate. They have no 'null' phases when tested with stationary gratings, and reveal their non-linearity by giving responses to gratings of some spatial phases that are composed partly or wholly of even harmonics of the stimulus frequency ('on-off' responses). 4. We compared simple receptive fields with their sensitivity to sinusoidal gratings of different spatial frequencies. Qualitatively, the most sensitive subregions of simple cells' receptive fields are roughly the same width as the individual bars of the gratings to which they are most sensitive. Quantitatively, their receptive field profiles measured with thin stationary lines, agree well with predicted profiles derived by Fourier synthesis of their spatial frequency tuning curves.

Action Potentials↗

Receptive field organization of complex cells in the cat's striate cortex.

1. All complex cells in the cat's striate cortex exhibit gross non-linearities of spatial summation when tested with sinusoidal grating stimuli. Their responses to moving gratings of all but the lowest spatial frequencies are usually dominated by a component that is not modulated by the passage of the bars of the grating across the receptive field. They give responses to temporally modulated stationary gratings that consist mostly of even harmonics of the stimulus frequency and that vary little in amplitude or wave form as the spatial phase of the grating is varied. 2. We compared complex cells' receptive fields with their sensitivity to sinusoidal gratings of different spatial frequencies. Qualitatively, the receptive fields are usually two to five times wider than the bars of the gratings that stimulate them most effectively. Quantitatively, the receptive field profiles of complex cells are invariably broader than those predicted by Fourier synthesis of their spatial frequency tuning curves, and in particular lack predicted spatially antagonistic regions. 3. We further examined the receptive field organization of these cells, using pairs of stationary lines flashed synchronously on their receptive fields. If both lines are of the same polarity (bright or dark), complex cells respond to the paired stimulus much less well than they do to either of its component bars, unless the bars are separated by less than about one quarter of the width of the receptive field. If the lines are of opposite polarity, one bright and one dark, the opposite situation obtains: closely spaced bars elicit small responses, while paired bars of larger separation are much more effective. In either case, the results are independent in general character of the absolute positions of the stimuli within the receptive field; rather, they depend in a manner characteristic of each cell on the relative positions of the two bars. 4. The two-line interaction profile that plots the change in a complex cell's response to one bar as a function of the position of a second added bar corresponds closely to the receptive field profile predicted from Fourier synthesis of the cell's spatial frequency tuning curve. These profiles may thus reveal the spatial characteristics of subunits within complex cell-receptive fields. We examined the nature of the interaction between these subunits by performing several two-line interaction experiments in which the onset of the second bar was delayed some time after the onset of the first. The results suggest that neighbouring subunits interact in a facilitatory fashion: for an interval after the presentation of one bar, responses to neighbouring bars are enhanced. 5. The subunits of a complex receptive field may, by their spatial properties, determine the spatial selectivities of complex cells, while the nature of the interaction among the subunits may determine these cells' sensitivity and selectivity for moving visual stimuli...

Action Potentials↗

Spatial and temporal contrast sensitivity of neurones in areas 17 and 18 of the cat's visual cortex.

1. We have examined the spatial and temporal tuning properties of 238 cortical neurones, recorded using conventional techniques from acutely prepared anaesthetized cats. We determined spatial and temporal frequency tuning curves using sinusoidal grating stimuli presented to each neurone's receptive field by a digital computer on a cathode ray tube. 2. We measured tuning curves either by determining response amplitude as a function of spatial or temporal frequency, or by measuring contrast sensitivity (the inverse of the contrast of the grating that just elicited a detectable response). The two measures give very similar tuning curves in all cases. 3. We recorded from 184 neurones in area 17; of these 156 had receptive fields within 5 degrees of the area centralis. The range of preferred spatial frequency for these neurones was 0.3--3 c/deg, and their spatial frequency tuning band widths varied from 0.7 to 3.2 octaves at half-amplitude. The most common band width was roughly 1.3 octaves. Simple and complex cells in area 17 did not differ in their distributions of preferred spatial frequency, although complex cells were, on average, slightly less selective for spatial frequency than simple cells. 4. We recorded from fifty-four neurones from area 18, and performed several experiments in which we recorded from corresponding portions of both area 17 and area 18 in the same electrode penetration. Neurones in area 18 preferred spatial frequencies that were, on average, one third as high as those preferred by area 17 neurones at the same retinal eccentricity. Thus the range of preferred spatial frequency in area eighteen cells having receptive fields within 5 deg of the area centralis was between less than 0.1 and 0.5 c/deg. The distributions of optimum spatial frequency in the two areas were practically non-overlapping at eccentricities as high as 15 deg, the greatest eccentricity we examined. Neurones in area 18 were about as selective for spatial frequency as were neurones in area 17. 5. We determined temporal frequency tuning characteristics for some neurones from each area, using gratings that moved steadily across the screen. Neurones from area 17 all responded well to low temporal frequencies, and less well to higher frequencies (in excess of, usually, 2 or 4 Hz). In contrast, neurones recorded from area 18 sometimes had similar tuning properties, but more commonly showed a pronounced reduction in response as the temporal frequency was moved either above or below some optimum value (usually 2--8 Hz). 6. We conclude from these results that areas 17 and 18 act in parallel to process different aspects of the visual information relayed from the retina via the lateral geniculate complex. Some or all of the differences between the areas may be attributable to the predominance of Y cell input to area 18 and the predominance of X cell input to area 17...

Action Potentials↗

Trichromatic colour opponency in ganglion cells of the rhesus monkey retina.

Two hundred and eleven colour-opponent ganglion cells were studied in the central 10 degrees of the retina of the rhesus monkey, to determine the inputs which they were receiving from different cone mechanisms. Spectral-sensitivity measurements in the presence of neutral and coloured back-grounds showed that 24% of these cells appeared to receive input from all three cone mechanisms. 2. In 3% of the cells, the red-sensitive cone mechanism opposed the blue- and green-sensitive ones. In 18% of the cells, the blue-sensitive cone mechanism opposed the green- and red-sensitive ones. In 3% of the cells, the green-sensitive cone mechanism opposed the blue- and red-sensitive ones. 3. In 12% of the cells receiving opponent green- and red-sensitive cone inputs, responses from the beta-band of the red-sensitive cone mechanism could be detected and distinguished from blue-sensitive cone input. 4. All cells receiving blue-sensitive cone input appeared to be trichromatic. The retinal distribution of cells with trichromatic input and that of cells with beta-band responses seemed to parallel the availability of blue-sensitive cones in the retinal area being considered. 5. The results indicate that trichromatic interactions in the macaque visual system begin in the retina.

Adaptation, Ocular↗