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Neurons in striate cortex limit the spatial and temporal resolution for detecting disparity modulation.
Neurons in striate cortex limit the spatial and temporal resolution for detecting disparity modulation.
Stereopsis is the process of seeing depth constructed from binocular disparity. The human ability to perceive modulation of disparity over space (Tyler, 1974; Prince and Rogers, 1998; Banks et al., 2004a) and time (Norcia and Tyler, 1984) is surprisingly poor, compared with the ability to detect spatial and temporal modulation of luminance contrast. In order to examine the physiological basis of this poor spatial and temporal resolution of stereopsis, I quantified responses to disparity modulation in disparity selective V1 neurons from four awake behaving monkeys. To study the physiological basis of the spatial resolution of stereopsis, I characterized the three-dimensional structure of 55 V1 receptive fields (RF) using random dot stereograms in which disparity varied as a sinusoidal function of vertical position (“corrugations”). At low spatial frequencies, this produced a modulation in neuronal firing at the temporal frequency of the stimulus. As the spatial frequency increased, the modulation reduced. The mean response rate changed little, and was close to that produced by a uniform stimulus at the mean disparity of the corrugation. In 48/55 (91%) of the neurons, the modulation strength was a lowpass function of spatial frequency. These results suggest that the neurons have fronto-parallel planar receptive fields, no disparity-based surround inhibition and no selectivity for disparity gradients. This scheme predicts a relationship between RF size and the high frequency cutoff. Comparison with independent measurements of RF size was compatible with this. All of this behavior closely matches the binocular energy model, which functionally corresponds to cross-correlation: the disparity modulated activity of the binocular neuron measures the correlation between the filtered monocular images. To examine the physiological basis of the temporal resolution of stereopsis, I measured for 59 neurons the temporal frequency tuning with random dot stereograms in which disparity varied as a sinusoidal function of time. Temporal frequency tuning in response to disparity modulation was not correlated with temporal frequency tuning in response to contrast modulation, and had lower temporal frequency high cutoffs on average. The temporal frequency high cut for disparity modulation was negatively correlated with the response latency, the speed of the response onset and the temporal integration time (slope of the line relating response phase and temporal frequency). Binocular cross-correlation of the monocular images after bandpass filtering can explain all these results. Average peak temporal frequency in response to disparity modulation was 2Hz, similar to the values I found in four human observers (1.5-3Hz). The mean cutoff spatial frequency, 0.5 cpd, was similar to equivalent measures of decline in human psychophysical sensitivity for such depth corrugations as a function of frequency (Tyler, 1974; Prince and Rogers, 1998; Banks et al., 2004a). This suggests that the human temporal and spatial resolution for stereopsis is limited by selectivity of V1 neurons. For both, space and time, the lower resolution for disparity modulation than for contrast modulation can be explained by a single mechanism, binocular cross-correlation of the monocular images. The findings also represent a significant step towards understanding the process by which neurons solve the stereo correspondence problem (Julesz, 1971).
binocular disparity, stereopsis, macaque V1, disparity modulation, temporal resolution, spatial resolution, physiology, single unit, human psychophysics
Nienborg, Hendrikje
2005
English
Universitätsbibliothek der Ludwig-Maximilians-Universität München
Nienborg, Hendrikje (2005): Neurons in striate cortex limit the spatial and temporal resolution for detecting disparity modulation.. Dissertation, LMU München: Faculty of Medicine
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Abstract

Stereopsis is the process of seeing depth constructed from binocular disparity. The human ability to perceive modulation of disparity over space (Tyler, 1974; Prince and Rogers, 1998; Banks et al., 2004a) and time (Norcia and Tyler, 1984) is surprisingly poor, compared with the ability to detect spatial and temporal modulation of luminance contrast. In order to examine the physiological basis of this poor spatial and temporal resolution of stereopsis, I quantified responses to disparity modulation in disparity selective V1 neurons from four awake behaving monkeys. To study the physiological basis of the spatial resolution of stereopsis, I characterized the three-dimensional structure of 55 V1 receptive fields (RF) using random dot stereograms in which disparity varied as a sinusoidal function of vertical position (“corrugations”). At low spatial frequencies, this produced a modulation in neuronal firing at the temporal frequency of the stimulus. As the spatial frequency increased, the modulation reduced. The mean response rate changed little, and was close to that produced by a uniform stimulus at the mean disparity of the corrugation. In 48/55 (91%) of the neurons, the modulation strength was a lowpass function of spatial frequency. These results suggest that the neurons have fronto-parallel planar receptive fields, no disparity-based surround inhibition and no selectivity for disparity gradients. This scheme predicts a relationship between RF size and the high frequency cutoff. Comparison with independent measurements of RF size was compatible with this. All of this behavior closely matches the binocular energy model, which functionally corresponds to cross-correlation: the disparity modulated activity of the binocular neuron measures the correlation between the filtered monocular images. To examine the physiological basis of the temporal resolution of stereopsis, I measured for 59 neurons the temporal frequency tuning with random dot stereograms in which disparity varied as a sinusoidal function of time. Temporal frequency tuning in response to disparity modulation was not correlated with temporal frequency tuning in response to contrast modulation, and had lower temporal frequency high cutoffs on average. The temporal frequency high cut for disparity modulation was negatively correlated with the response latency, the speed of the response onset and the temporal integration time (slope of the line relating response phase and temporal frequency). Binocular cross-correlation of the monocular images after bandpass filtering can explain all these results. Average peak temporal frequency in response to disparity modulation was 2Hz, similar to the values I found in four human observers (1.5-3Hz). The mean cutoff spatial frequency, 0.5 cpd, was similar to equivalent measures of decline in human psychophysical sensitivity for such depth corrugations as a function of frequency (Tyler, 1974; Prince and Rogers, 1998; Banks et al., 2004a). This suggests that the human temporal and spatial resolution for stereopsis is limited by selectivity of V1 neurons. For both, space and time, the lower resolution for disparity modulation than for contrast modulation can be explained by a single mechanism, binocular cross-correlation of the monocular images. The findings also represent a significant step towards understanding the process by which neurons solve the stereo correspondence problem (Julesz, 1971).