In age-related macular degeneration (AMD), the processing of fine details in a visual scene, based on a high spatial frequency processing, is impaired, while the processing of global shapes, based on a low spatial frequency processing, is relatively well preserved. The present fMRI study aimed to investigate the residual abilities and functional brain changes of spatial frequency processing in visual scenes in AMD patients. AMD patients and normally sighted elderly participants performed a categorization task using large black and white photographs of scenes (indoors vs. outdoors) filtered in low and high spatial frequencies, and nonfiltered. The study also explored the effect of luminance contrast on the processing of high spatial frequencies. The contrast across scenes was either unmodified or equalized using a root-mean-square contrast normalization in order to increase contrast in high-pass filtered scenes. Performance was lower for high-pass filtered scenes than for low-pass and nonfiltered scenes, for both AMD patients and controls. The deficit for processing high spatial frequencies was more pronounced in AMD patients than in controls and was associated with lower activity for patients than controls not only in the occipital areas dedicated to central and peripheral visual fields but also in a distant cerebral region specialized for scene perception, the parahippocampal place area. Increasing the contrast improved the processing of high spatial frequency content and spurred activation of the occipital cortex for AMD patients. These findings may lead to new perspectives for rehabilitation procedures for AMD patients.

We measured human frequency response functions for eleven angular frequency filters using a forced-choice procedure in a supra-threshold summation paradigm. Each of the eleven functions of 17 experimental conditions was measured 4-9 times among 12 observers. Results show that, for the arbitrarily selected filter phases, maximum summation effect occurred at test frequency for all filters. These results lead to the conclusion that there are narrow-band angular frequency filters operating in human visual system mostly through summation surrounded by inhibition at the specific test frequency ranges. Our previous suggestion (Simas & Santos, 2002), arguing that summation for the higher angular frequency filters should occur if background angular frequency contrast were set to a maximum of 5 times the test frequency threshold, was supported.

The role of different spatial frequency bands on face gender and expression categorization was studied in three experiments. Accuracy and reaction time were measured for unfiltered, low-pass (cut-off frequency of 1 cycle/deg) and high-pass (cut-off frequency of 3 cycles/deg) filtered faces. Filtered and unfiltered faces were equated in root-mean-squared contrast. For low-pass filtered faces reaction times were higher than unfiltered and high-pass filtered faces in both categorization tasks. In the expression task, these results were obtained with expressive faces presented in isolation (Experiment 1) and also with neutral-expressive dynamic sequences where each expressive face was preceded by a briefly presented neutral version of the same face (Experiment 2). For high-pass filtered faces different effects were observed on gender and expression categorization. While both speed and accuracy of gender categorization were reduced comparing to unfiltered faces, the efficiency of expression classification remained similar. Finally, we found no differences between expressive and non expressive faces in the effects of spatial frequency filtering on gender categorization (Experiment 3). These results show a common role of information from the high spatial frequency band in the categorization of face gender and expression.

The responses of simple cells (recorded from within the striate visual cortex) were measured as a function of the contrast and the frequency of sine-wave grating patterns in order to explore the effect of contrast on the spatial and temporal phase transfer functions and on the spatiotemporal receptive field. In general, as the contrast increased, the phase of the response advanced by approximately 45 ms (approximately one-quarter of a cycle for frequencies near 5 Hz), although the exact value varied from cell to cell. The dynamics of this phase-advance were similar to the dynamics of the amplitude: the amplitude and the phase increased in an accelerating fashion at lower contrasts and then saturated at higher contrasts. Further, the gain for both the amplitude and the phase appeared to be governed by the magnitude of the contrast rather than the magnitude of the response. For the spatial phase transfer function, variations in contrast had little or no systematic effect; all of the phase responses clustered around a single straight line, with a common slope and intercept. This implies that the phase-advance was not due to a change in the spatial properties of the neuron; it also implies that the phase-advance was not systematically related to the magnitude of the response amplitude. On the other hand, for the temporal phase transfer function, the phase responses fell on five straight lines, related to the five steps in contrast. As the contrast increased, the phase responses advanced such that both the slope and the intercept were affected. This implies that the phase-advance was a result of contrast-induced changes in both the response latency and the shape/symmetry of the temporal receptive field.

White noise stimuli were used to estimate second-order kernels for complex cells in cortical area VI of the macaque monkey, and drifting grating stimuli were presented to the same sample of neurons to obtain orientation and spatial-frequency tuning curves. Using these data, we quantified how well second-order kernels predict the normalized tuning of the average response of complex cells to drifting gratings.

The estimated second-order kernel of each complex cell was transformed into an interaction function defined over all spatial and temporal lags without regard to absolute position or delay. The Fourier transform of each interaction function was then computed to obtain an interaction spectrum. For a cell that is well modeled by a second-order system, the cell’s interaction spectrum is proportional to the tuning of its average spike rate to drifting gratings. This result was used to obtain spatial-frequency and orientation tuning predictions for each cell based on its second-order kernel. From the spatial-frequency and orientation tuning curves, we computed peaks and bandwidths, and an index for directional selectivity.

We found that the predictions derived from second-order kernels provide an accurate description of the change in the average spike rate of complex cells to single drifting sine–wave gratings. These findings are consistent with a model for complex cells that has a quadratic spectral energy operator at its core but are inconsistent with a spectral amplitude model.

Measurements were made of the optimal velocity for drifting bar-shaped stimuli to excite striate cortex neurons of the cat. These data were compared to the optimal spatial and temporal frequencies of the same neurons, as determined with drifting sine-wave grating stimuli. A systematic relationship was revealed, whereby those neurons preferring higher velocities of bar motion also preferred lower spatial and higher temporal frequencies of gratings. The optimal bar velocity for a given neuron could be quantitatively predicted from the ratio of that neuron's optimal temporal frequency to its optimal spatial frequency.

It is believed that spatial summation in most simple cells is a linear process. If this were so, then the Fourier transform of a simple cell's line weighting function should predict the cell's spatial frequency tuning curve. We have compared such predictions with experimental measurements and have found a consistent discrepancy: the predicted tuning curve is much too broad. We show qualitatively that this kind of discrepancy is consistent with the well-known threshold nonlinearity shown by most cortical cells. We have tested quantitatively whether a response threshold could explain the observed disagreements between predictions and measurements: a least-squares minimization routine was used to fit the inverse Fourier Transform of the measured frequency tuning curve to the measured line weighting function. The fitting procedure permitted us to introduce a threshold to the reconstructed line weighting function. The results of the analysis show that, for all of the cells tested, the Fourier transforms produced better predictions when a response threshold was included in the model. For some cells, the actual magnitude of the response threshold was measured independently and found to be compatible with that suggested by the model. The effects of nonlinearities of spatial summation are considered.

Averaged grating-evoked cortical potentials were recorded from area 17 of awake cats. Peak latency of early components of the visual-evoked potential (VEP) response to stimulus onset increased as a function of spatial frequency, while amplitude tended to be largest at intermediate spatial frequencies. Latency increased and amplitude generally decreased to lower spatial-frequency stimuli (<0.25 cycle/deg) in the presence of a uniform flickering field (UFF). The UFF had a relatively small or opposite effect on peak latency and amplitude for higher spatial-frequency stimuli (>0.50 cycle/deg). The VEP response to stimulus offset was present only at low spatial frequencies and was virtually eliminated by the presence of the UFF. The effects were similar whether the target and UFF background were simultaneously presented or briefly separated; however, the UFF had no effect when the two were spatially separated. The effects of the UFF background on VEP onset response increased with increasing temporal frequency from 2–8 Hz; offset responses were affected similarly at all temporal frequencies. These effects are similar to those observed in humans and suggest that two spatio-temporally tuned mechanisms contribute to the early VEP response. In the cat, the mechanisms seem to correspond to X and Y cells in the dorsal lateral geniculate nucleus.

The amplitude spectra of simple cells in areas 17 and 18 were estimated in two and three dimensions (2–D and 3–D) using drifting sinusoidal gratings. In 2–D, responses were sampled with 16 x 16 resolution in spatial and temporal frequency at the optimal orientation. In 3–D, responses were sampled with 12 x 12 x 10 resolution in spatial frequency, orientation, and temporal frequency. For 45/50 cells studied, the spatial attributes of the receptive fields (RFs) were independent of temporal frequency except for a scale factor. The five exceptions to this general finding could be described as follows: For four area 17 cells, responses in the null direction increased with temporal frequency, reducing direction selectivity. For one area 18 cell, the optimal spatial frequency increased with temporal frequency and vice versa. The 2–D discrete Fourier transform was applied to all of the estimated amplitude spectra assuming zero spatial and temporal phase. These transforms were compared with the results of first-order reverse correlations as described in the previous paper (McLean et al., 1994). Direction selective cells exhibited excitatory subregions that were obliquely oriented in space-time in both the raw correlation data and inverse transforms of the spectral data. The slopes of the subregions found in these two measures were highly correlated. Direction indices obtained from space and frequency domain measures were comparable. We demonstrate that the spectral response profiles of most simple cells are aligned with the coordinate axes in frequency domain. That is, they may be considered one-quadrant separable, suggesting that these cells are not velocity tuned per se, but are tuned for spatiotemporal frequency. The spectral bandwidth establishes the range of velocities to which these cells will respond. These findings are consistent with the one-quadrant separability constraint of linear quadrature models. We conclude that most simple cells perform as roughly linear filters in two dimensions of space and time.

The existence in cat cortex of linearly operating simple cells with relative spatial-frequency bandwidths of between 0.5 and 1 octave is confirmed.

The superior colliculus (SC) projects to all layers of the cat's lateral geniculate nucleus (LGN) and thus is in a position to influence information transmission through the LGN. We investigated the function of the tecto-geniculate pathway by studying the responses of cat LGN neurons before, during, and after inactivating the SC with microinjections of lidocaine. The LGN cells were stimulated with drifting sine-wave gratings that varied in spatial frequency and contrast. Among 71 LGN neurons that were studied, 53 showed a statistically significant change in response during SC inactivation. Control experiments with mock injections indicated that some changes could be attributed to slow waxing and waning of responsiveness over time. However, this could not account for all of the effects of SC inactivation that were observed. Forty cells showed changes that were attributed to the removal of tecto-geniculate influences. About equal numbers of cells showed increases (22 cells) and decreases (18 cells) in some aspect of their response to visual stimuli during SC inactivation. The proportion of cells that showed tecto-geniculate influences was somewhat higher in the C layers (68% of the cells) than in the A layers (44% of the cells). In addition, among cells that showed a significant change in maximal response to visual stimulation, the change was larger for cells in the C layers (64% average change) than in the A layers (26% average change) and it was larger for W cells (61% average change) than for X and Y cells (29% average change). Nearly all of the X cells that showed changes had an increase in response, and nearly all of the Y cells had a decrease in response. In addition, across all cell classes, 80% of the cells with receptive fields < 15 deg from the area centralis had an increase in response, and 80% of the cells with receptive fields > 15 deg from the area centralis had a decrease in response. None of the LGN cells had significant changes in spatial resolution, and only three cells had changes in optimal spatial frequency. Ten cells had a change in contrast threshold, 25 cells had a change in contrast gain, and 29 cells had a change in the maximal response to a high-contrast stimulus. Thus, our results suggest that the tecto-geniculate pathway has little or no effect on spatial processing by LGN neurons. Rather, the major influence is on maximal response levels and the relationship between response and stimulus contrast. Several hypotheses about the role of the tecto-geniculate pathway in visual behavior are considered.

Age-related macular degeneration (AMD) is a major cause of visual impairment in people older than 50 years in Western countries, affecting essential tasks such as reading and face recognition. Here we investigated the mechanisms underlying the deficit in recognition of facial expressions in an AMD population with low vision. Pictures of faces displaying different emotions with the mouth open or closed were centrally displayed for 300 ms. Participants with AMD with low acuity (mean 20/200) and normally sighted age-matched controls performed one of two emotion tasks: detecting whether a face had an expression or not (expressive/non expressive (EXNEX) task) or categorizing the facial emotion as happy, angry, or neutral (categorization of expression (CATEX) task). Previous research has shown that healthy observers are mainly using high spatial frequencies in an EXNEX task while performance at a CATEX task was preferentially based on low spatial frequencies. Due to impaired processing of high spatial frequencies in central vision, we expected and observed that AMD participants failed at deciding whether a face was expressive or not but categorized normally the emotion of the face (e.g., happy, angry, neutral). Moreover, we observed that AMD participants mostly identified emotions using the lower part of the face (mouth). Accuracy did not differ between the two tasks for normally sighted observers. The results indicate that AMD participants are able to identify facial emotion but must base their decision mainly on the low spatial frequencies, as they lack the perception of finer details.

The aim of this study was to use non-invasive magneto-encephalographic techniques, together with visual stimulus paradigms that can psychophysically separate the M- and P-pathways, to examine the physiological relations of the pathways at the fovea with (1) the magneto-encephalography components M70 and M100 (in latency and amplitude), and (2) the cortical oscillatory activities (alpha, beta, and gamma), respectively. The checkerboard stimuli accompanied with different spatial frequencies (SFs) (0.5 or 4 cycles per degree) were presented (within 2° of the retinal center) to six healthy subjects by using steady-pedestal and pulse paradigms, which could activate distinct populations of M- and P-neurons. SF analyzed brain responses in each paradigm. The results show a consistent trend in M70 and M100 with increased latencies and amplitudes in response to the high SF. Mean while, the beta to gamma activities are apparently enhanced by the stimulus of high SF, especially under pulse paradigm (p = 0.03). In this study, we suggest that M70 can be a good clue to characterize the P-pathway. Moreover, in the frequency analysis, the beta oscillations may serve for more detailed visual information, while the gamma oscillations seem to reflect the signal processing in the P-pathway and with sensitivity to the fovea.

The retinal dopaminergic system is a global regulator of retinal function. Apart from the fact that the rates of dopamine synthesis and release are increased by increasing illumination, the visual image parameters that influence dopaminergic function are mostly unknown. Roles for spatial and temporal frequency and image contrast are suggested by the effects of form-deprivation with a diffusing goggle. Form-deprivation reduces the rates of dopamine synthesis and release, and induces myopia, which is prevented by dopamine agonists. Our purpose here was to identify visual stimulus parameters that activate dopaminergic amacrine cells and elicit dopamine release. White Leghorn cockerels 4–7 days old were exposed to 2 h of form-deprivation, reduced light intensity, or stimuli of varied temporal or spatial frequency. Activation of dopaminergic neurons, labeled for tyrosine hydroxylase (TH), was assessed with immunocytochemistry for c-Fos, and dopamine release was measured by HPLC analysis of dopamine metabolite accumulation in the vitreous body. Form-deprivation did not reduce TH+ cell activation or vitreal dopamine metabolite accumulation any more than did neutral-density filters of approximately equal transmittance. TH+ cell activation and vitreal metabolite accumulation were not affected significantly by exposure to 2, 5, 10, 15, or 20 Hz stroboscopic stimulation on a dark background, or by sine-wave gratings of 0.089, 0.44, 0.89, 1.04, or 3.13 cycles/deg compared to a uniform gray target of equal mean luminance. These data indicate that the retinal dopaminergic system does not respond readily to short-term changes in visual stimulus parameters, other than light intensity, under the conditions of these experiments.

The mosaics of S-cones and the neurons to which they are connected are relatively well characterized, so the S-cone system is a good vehicle for exploring how the sampling of the retinal image controls visual performance. We used an interferometer to measure the grating acuity of the S-cone system in the fovea and at a range of eccentricities out to 20 deg. We also developed a simple model observer that, by assuming only that cone pathways are noisy and that signals are subject to eccentricity-dependent postreceptoral pooling, predicts the measured acuities from the sampling properties of the S-cone mosaic. The amount of pooling required to explain performance is consistent with that suggested by anatomical and physiological measurements.

We report results of numerical simulations for a model of generation of orientation selectivity in macaque striate cortex. In contrast to previous models, where the initial orientation bias is generated by convergent geniculate input to simple cells and subsequently sharpened by lateral circuits, our approach is based on anisotropic intracortical excitatory connections which provide both the initial orientation bias and its subsequent amplification. Our study shows that the emerging response properties are similar to the response properties that are observed experimentally, hence the hypothesis of an intracortical generation of orientation bias is a sensible alternative to the notion of an afferent bias by convergent geniculocortical projection patterns. In contrast to models based on an afferent orientation bias, however, the “intracortical hypothesis” predicts that orientation tuning gradually evolves from an initially nonoriented response and a complete loss of orientation tuning when the recurrent excitation is blocked, but new experiments must be designed to unambiguously decide between both hypotheses.