Detection and identification of near-threshold visual patterns.

Temporal and spatial frequency tuning of the flicker motion aftereffect.

The visual system demonstrates significant differences in information processing abilities between the central and peripheral parts of the visual field. Optical imaging based on intrinsic signals was used to investigate the difference in stimulus spatial and temporal frequency interactions related to receptive field eccentricity in the cat area 18. Changing either the spatial or the temporal frequency of grating stimuli had a significant impact on responses in the cortical areas corresponding to the centre of the visual field and more peripheral parts at 10 degrees eccentricity. The cortical region corresponding to the centre of the gaze was tuned to 0.4 cycles per degree (c/deg) for spatial frequency and 2Hz for temporal frequency. In contrast, the cortical region corresponding to the periphery of the visual field was tuned to a lower spatial frequency of 0.15c/deg and a higher temporal frequency of 4Hz. Interestingly, when we simultaneously changed both the spatial frequency and the temporal frequency of the grating stimuli, the responses were significantly different from those estimated with an assumption of independence between the spatial and temporal frequency in the cortical region corresponding to the periphery of the visual field. However, in the cortical area corresponding to the centre of the gaze, spatial frequency showed significant independence from temporal frequency. These properties support the notion of relative specialization of visual information processing for peripheral representations in cortical areas.

Figure-ground organization of an ambiguous pattern can be manipulated by the spatial and temporal frequency content of the two regions of the pattern. Controlling for space-averaged luminance and perceived contrast, we tested patterns in which the two regions of the ambiguous pattern contained sine-wave gratings of 8, 4, 1, or 0.5 cycles per degree (cpd) undergoing on:off flicker at the rates of 0, 3.75, 7.5, or 15 Hz. For a full set of combinations of temporal frequency differences, with each spatial frequency the higher temporal frequency was seen as background for more of the viewing time. For two spatial frequency combinations, 1 and 4 cpd, and 1 and 8 cpd, tested under each of the four temporal frequencies, the lower spatial frequency region was seen as the background for more of the viewing time. When the effects of spatial and temporal frequency were set in opposition, neither was predominant in determining perceptual organization. It is suggested that figure-ground organization may parallel the sustained-transient response characteristics of the visual system.

The red-green axis of colour vision evolved recently in primate evolutionary history. Signals serving red-green colour vision travel together with signals serving spatial vision, in the parvocellular (PC) division of the subcortical visual pathway. However, the question of whether receptive fields of PC pathway cells are specialized to transmit red-green colour signals remains unresolved. We addressed this question in single-cell recordings from the lateral geniculate nucleus of anaesthetized marmosets. Marmosets show a high proportion of dichromatic (red-green colour-blind) individuals, allowing spatial and colour tuning properties of PC cells to be directly compared in dichromatic and trichromatic visual systems. We measured spatial frequency tuning for sine gratings that provided selective stimulation of individual photoreceptor types. We found that in trichromatic marmosets, the foveal visual field representation is dominated by red-green colour-selective PC cells. Colour selectivity of PC cells is reduced at greater eccentricities, but cone inputs to centre and surround are biased to create more selectivity than predicted by a purely 'random wiring' model. Thus, one-to-one connections in the fovea are sufficient, but not necessary, to create colour-selective responses. The distribution of spatial tuning properties for achromatic stimuli shows almost complete overlap between PC cells recorded in dichromatic and trichromatic marmosets. These data indicate that transmission of red-green colour signals has been enabled by centre-surround receptive fields of PC cells, and has not altered the capacity of PC cells to serve high-acuity vision at high stimulus contrast.

Spatiotemporal Frequency and Direction Sensitivities of Human Visual Areas Measured Using fMRI

The present experiments examined the extent to which binocular processing in the cat's dorsal lateral geniculate nucleus (LGN) depends upon the spatial frequency, orientation, and direction of movement of stimuli presented to the nondominant eye. In Experiment 1, we tested the effects of varying these stimulus parameters on the responses of LGN cells to nondominant-eye stimulation. Sixteen of 34 cells tested had statistically significant responses to the nondominant eye and, in agreement with a previous study (Guido et al. 1989), the responsive cells were spatial-frequency sensitive. However, there was little evidence for orientation or direction sensitivity in responses to the nondominant eye: changes in discharge with changes in stimulus orientation and direction were small and were statistically significant in only nine of the cells. In Experiment 2, we tested the effects of varying spatial frequency, orientation, and direction of movement of stimuli presented to the nondominant eye on its ability to influence responses to the dominant eye (i.e., on binocular interactions). The dominant eye was stimulated with the optimal spatial frequency for the cell being tested. For 22 of 45 cells tested, nondominant-eye stimulation had a statistically significant effect on the response to the dominant eye. Fourteen of these cells showed band-pass spatial-frequency sensitivity in the nondominant-eye influence, and eight showed low-pass spatial-frequency sensitivity. However, only 11 of the cells had statistically significant variations in their binocular interactions that depended on the orientation or direction of stimuli presented to the nondominant eye. Furthermore, even for those cells, the effect of varying orientation and direction was only about half as strong as the effect of varying spatial frequency. We conclude that binocular processing in the LGN, including responses to the nondominant eye and nondominant-eye influences on responses to the dominant eye, are affected significantly by the spatial frequency of the nondominant-eye stimulus and relatively little by stimulus orientation or direction of movement. The significance of these findings for understanding the functions of LGN binocular processing is discussed.

The receptive field of a visual neuron is the particular region in which a stimulus will modify the firing of that neuron. Retinal ganglion cells have a coaxial-shaped structure formed by the central excitatory and outer inhibitory regions. These regions respond oppositely to light. This shape can be mathematically modeled by a balanced Difference of Gaussian (DOG) function. In digital image processing, the DOG filter is used for detecting objects' edges. Bacteriorhodopsin (bR) is a photosensitive protein which resembles the visual pigment rhodopsin. When bR is illuminated, it generates a positive transient photocurrent, and, when light is switched off, a negative transient photocurrent is produced. This peculiar behavior is similar to the response of ganglion receptive fields. In this study, we fabricate a two-dimensional binarized DOG (b-DOG) filter, in which the central part mimics the excitatory part of the ganglion cell receptive fields, whereas the outer ring reproduces the effect observed in their outer inhibitory regions. This filter consists of photosensitive protein bR films and electrolyte solution that are sandwiched between ITO electrodes. To analyze the spatial-temporal frequency sensitivity, we use moving sine wave gratings with different pitches with a controlled scanning speed. When the temporal frequency is kept constant, the spatial frequency sensitivity matched with the Fourier transform of the b-DOG function. On the other hand, when the spatial frequency does not vary, the temporal frequency sensitivity corresponded well with the Fourier transform of Difference of Gamma function. Difference of Gamma function is known as the impulse response of the visual nerve of animals. We separately analyze the independent spatial and temporal frequencies collected from the spatial-temporal characteristics of the filter. The spatial-temporal frequency characteristics of b-DOG filter are similar to those of the X-type retinal ganglion cells. However, unlike the retinal ganglion cell, the b-DOG filter is a perfect linear filter. The analog image processing using this b-DOG filter is performed by scanning a standard test image. It is found that an edge can be detected just by scanning the image and plotting the zero-crossing point. With increasing input image size, the spatial frequency peak detected by the b-DOG filter shifts towards higher frequencies. Since there are only sharp edges where high spatial frequency components are present, the detected results of the analog filter are similar to that of the digital filter. Consequently, the phase difference between the input and output image is approximately the same for all pixels, and it agrees with the digital edge detection results. Some illusion images are scanned on the b-DOG filter to verify the occurrence of the illusion similar to vision. The Hermann grid illusion generated with lower-order vision was observed. It is found that no luminance information is necessary for the Herman grid illusion, because this b-DOG filter does not extract image luminance. The visual function elements similar to X-type retinal ganglion cells prepared in this study are useful to constructively understand the visual information processing mechanism of the organism.

Tuning for speed is one key feature of motion-selective neurons in the middle temporal visual area of the macaque cortex (MT, or V5). The present paper asks whether speed is coded in a way that is invariant to the shape of the moving stimulus, and if so, how. When tested with single sine-wave gratings of different spatial and temporal frequencies, MT neurons show a continuum in the degree to which preferred speed depends on spatial frequency. There is some dependence in 75% of MT neurons, and the other 25% maintain speed tuning despite changes in spatial frequency. When tested with stimuli constructed by adding two superimposed sine-wave gratings, the preferred speed of MT neurons becomes less dependent on spatial frequency. Analysis of these responses reveals a speed-tuning nonlinearity that selectively enhances the responses of the neuron when multiple spatial frequencies are present and moving at the same speed. Consistent with the presence of the nonlinearity, MT neurons show speed tuning that is close to form-invariant when the moving stimuli comprise square-wave gratings, which contain multiple spatial frequencies moving at the same speed. We conclude that the neural circuitry in and before MT makes no explicit attempt to render MT neurons speed-tuned for sine-wave gratings, which do not occur in natural scenes. Instead, MT neurons derive form-invariant speed tuning in a way that takes advantage of the multiple spatial frequencies that comprise moving objects in natural scenes.

According to a modern view, simple cells of the cat striate cortex are considered to operate as a part of Fourier analysis system thus leading to the idea that the operational mechanism of the visual cortex is concerned with the analysis of spatial frequencies. Nevertheless if simple cells are really concerned only with the analysis of spatial frequencies there should exist a strict relationship between their spatial frequency selectivity and the spatial organization of their receptive fields. This is because it is the spatial organization of the spatial frequency detector i.e. the cell's receptive field that determines the cell's spatial frequency selectivity. Since the quantitative analysis of the spatial organization of simple cell receptive fields has shown that the receptive fields of these cells are made up of two to four antagonistic subregions it follows that striate simple cells are concerned with the analysis of only a selected range of spatial frequencies. Therefore striate simple cells must be considered analysers of visual signals both in spatial frequencies as well as in spatial positions.

1. The spatial and temporal response properties of single cells in area 21a of the anaesthetized cat were assessed using drifting sinusoidal gratings presented at the optimum orientation for each cell. 2. Responses to sinusoidal gratings were dominated by an elevation of the mean discharge, with a relatively small modulated component at the temporal frequency of grating drift. The relative modulation ratio for the majority of cells was less than 1, similar to complex cells in the striate cortex. 3. Of those cells responsive to stimulation with sinusoidal gratings, 94% displayed spatial bandpass characteristics. Values derived from spatial frequency tuning curves were: mean optimum spatial frequency, 0.26 cycles deg-1; mean spatial resolution, 0.86 cycles deg-1; mean spatial bandwidth, 1.8 octaves; and mean normalized bandwidth, 1.3. Two cells (6%) displayed spatial low-pass characteristics. 4. Approximately half our sample of cells (44%) displayed temporal low-pass tuning, while 35% displayed temporal bandpass characteristics. The mean optimum temporal frequency of bandpass cells was 3.3 Hz and the mean temporal bandwidth 1.9 octaves. The remaining cells were classified as temporal broadband (17%) and temporal high-pass (4%). 5. We conclude that the dominant functional input to cells with relatively high spatial frequency selectivity and/or temporal low-pass response properties most probably arises from area 17. The responses of the remaining cells may be explained by input from area 17 or 18.

In addition to a topographic map of the retina, mammalian visual cortex contains superimposed, orderly periodic maps of features such as orientation, eye dominance, direction of motion and spatial frequency. There is evidence that these maps are overlaid so as to ensure that all combinations of the different parameters are represented as uniformly as possible across visual space. However, it is unknown to what extent geometrical factors limit the number of periodic maps which might simultaneously be present, given this constraint. This paper attempts to investigate the question by using a dimension reduction model to generate maps of simple, many- dimensional feature spaces onto a model two-dimensional cortex. The feature space included a model retina, plus N binary variables, corresponding to parameters such as ocular dominance or spatial frequency. The results suggest that geometrical factors do not sharply limit the ability of the cortex to represent combinations of parameters in spatially superimposed maps of similar periodicity. Considerations of uniform coverage suggest an upper limit of six or seven maps. A higher limit, of about nine or ten, may be imposed by the numbers of neurons (or minicolumns) available to represent each of 2(N) features within a given small region of cortex.

Thresholds for detecting the direction of motion of drifting (8-Hz) vertical gratings [of spatial frequencies 0.1, 1.0, and 10.0 cycles per degree (c/deg)] were measured in the presence of masks that varied in both spatial frequency and orientation. Masks with different temporal properties were used. The specificity of masking was also measured for a stationary test grating of spatial frequency 3.0 c/deg. After suitable scaling and transformation, the masking data gave an estimate of the two-dimensional spatial-frequency tuning surface of cortical detector units in human vision. With the assumption of small-signal linearity and zero phase, the tuning surfaces were inverse Fourier transformed to give an indication of the size and structure of the psychophysical receptive fields of detector units. The results obtained with drifting test gratings and jittering (random phase) mask gratings indicate that motion-detector receptive fields increase in size (in cycles) with increasing spatial frequency but, at all spatial scales, have a length-width ratio of 1. These results are in close agreement with the summation results reported in J. Opt. Soc. Am. A 8, 1330 (1991). Using the same jittering mask stimuli and stationary test gratings, we confirm reports by Daugman [Vision Res. 24, 891 (1984)] and Harvey and Doan [J. Opt. Soc. Am. A 7, 116 (1990)] that motion-independent units have elongated receptive fields with a length-width ratio near 1.8. We conclude that the receptive fields of motion-dependent and -independent mechanisms in human vision are fundamentally different. The possibility that the orientation selectivity of a motion unit is sharpened by its selectivity for direction of motion is discussed.

1. We recorded local field potentials in the parafoveal representation in the primary visual cortex of anesthetized and paralyzed macaque monkeys with a multicontact electrode that provided for sampling of neural activity at 16 sites along a vertical penetration. Differential recordings at adjacent contacts were transformed into an estimate of current source density (CSD), to provide a measure of local neural activity. 2. We used m-sequence stimuli to map the region of visual space that provided input to the recording site. The local field potential recorded in macaque V1 has a population receptive field (PRF) size of approximately 2 deg2. 3. We assessed spatial tuning by the responses to two-dimensional Gaussian noise, spatially filtered to retain power only within one octave. Responses to achromatic band-limited noise stimuli revealed a prominent band-pass spatial tuning in the upper layers, but a more low-pass spatial tuning in lower layers. 4. We assessed orientation tuning by the responses to band-limited noise whose spectrum was further restricted to lie within 45 degrees wedges. The local field potential showed evidence of orientation tuning at most sites. Orientation tuning in upper and lower layers was manifest by systematic variations not only in response size but also in response dynamics. 5. We assessed chromatic tuning by the responses to isotropic band-limited noise modulated in a variety of directions in tristimulus space. Some lower-layer locations showed a nulling of response under near-isoluminant conditions. However, response dynamics in upper and lower layers depended not only on luminance contrast, but also on chromatic inputs. 6. Responses to near-isoluminant stimuli and to low-contrast luminance modulation were shifted to lower spatial frequencies. 7. We determined the extent to which various temporal frequencies in the response conveyed information concerning spatial frequency, orientation, and color under the steady-state conditions used in these studies. In each case, information is distributed in the response dynamics across a broad temporal frequency range, beginning at 4 Hz (the lowest frequency used). For spatial frequency the information rate remains significant up to at least 25 Hz. For orientation tuning and chromatic tuning, the information rate is lower overall and remains significant up to 13 Hz. In contrast, for texture discrimination, information is shifted to lower temporal frequencies.

Spatial frequency characteristics of nearby neurons in cats’ visual cortex

Objective To compare the difference between black-on-white pattern stimulation visual evoked potential (P-VEP) and blue-on-yellow pattern stimulation vision evoked potential in normal subjects.Method RETIport visual electrophysiography was used to record P-VEP at different spatial frequencies (60',30',15') and different color stimulations (black-on-white,blue-on-yellow)in 42 normal subjects (84 eyes).P100 wave latencies and N75～P100 wave amplitudes were compared.Results P100 wave latencies and N75～P100 wave amplitudes were statistics different (P ＜0.01) under different color stimulations (black-on-white,blue-on-yellow) at 60' spatial frequencies;N75～P100 wave amplitudes were statistics different (P ＜0.01) under different color stimulations (black-on-white,blue-on-yellow) at 30' and 15' spatial frequencies; P100 wave latencies were not statistics different (P ＞0.05) under different color stimulations (black-on-white,blue-on-yellow) at 30' and 15' spatial frequencies.Conclusions Blue-on-yellow P-VEP should establish its own normal value in clinic use,our research made useful information for clinic use. Key words: Color vision; Pattern visual evoked potential; Glaucoma