Forward and backward visual masking in schizophrenia: influence of age

BACKGROUND: Visual masking tasks assess the earliest stages of visual processing. This study was conducted to address: (1) whether schizophrenia patients show masking deficits after controlling for sensory input factors; (2) whether patients have relatively intact forward masking (when the mask precedes the target) compared with backward masking (when the mask follows the target); and (3) whether the masking deficits in schizophrenia reflect an accelerated age-related decline in performance. METHOD: A staircase method was used to ensure that the unmasked target identification was equivalent across subjects to eliminate any confounding due to differences in discrimination of simple perceptual inputs. Three computerized visual masking tasks were administered to 120 schizophrenia patients (ages 18-56) and 55 normal comparison subjects (ages 19-54) under both forward and backward masking conditions. The tasks included: (1) locating a target; (2) identifying a target with a high-energy mask; and (3) identifying a target with a low-energy mask. RESULTS: Patients showed deficits across all three masking tasks. Interactions of group by forward versus backward masking were not significant, suggesting that deficits in forward and backward masking were comparable. All three conditions showed an age-related decline in performance and rates of decline were comparable between patients and controls. Two of the masking conditions showed increased rates of decline in backward, compared to forward, masking. CONCLUSIONS: We found age-related decline in performance that was comparable for the two groups. In addition, we failed to find evidence of a relative sparing of forward masking in schizophrenia. These results suggest that: (1) early visual processing deficits in schizophrenia are not due to a simple perceptual input problem; (2) sustained channels are involved in the masking deficit (in addition to transient channels); and (3) for the age range in this study, these deficits in schizophrenia are not age-related.     

Transcranial magnetic stimulation in the visual system. I. The psychophysics of visual suppression

When applied over the occipital pole, transcranial magnetic stimulation (TMS) disrupts visual perception and induces phosphenes. Both the underlying mechanisms and the brain structures involved are still unclear. The first part of the study characterizes the suppressive effect of TMS by psychophysical methods. Luminance increment thresholds for orientation discrimination were determined in four subjects using an adaptive staircase procedure. Coil position was controlled with a stereotactic positioning device. Threshold values were modulated by TMS, reaching a maximum effect at a stimulus onset asynchrony (SOA) of approx. 100 ms after visual target presentation. Stronger TMS pulses increased the maximum threshold while decreasing the SOA producing the maximum effect. Slopes of the psychometric function were flattened with TMS masking by a factor of 2, compared to control experiments in the absence of TMS. No change in steepness was observed in experiments using a light flash as the mask instead of TMS. Together with the finding that at higher TMS intensities, threshold elevation occurs even with shorter SOAs, this suggests lasting inhibitory processes as masking mechanisms, contradicting the assumption that the phosphene as excitatory equivalent causes masking. In the companion contribution to this one we present perimetric measurements and phosphene forms as a function of the stimulation site in the brain and discuss the putative generator structures.     

Suppression on neuronal responses by a metacontrast masking stimulus in monkey V4

We studied the temporal characteristics of suppression in area V4 of the monkey using a visual stimulus for metacontrast masking. Visual responses of V4 neurons to a brief test stimulus presented within the receptive field were recorded, and the effect of a mask stimulus that did not spatially overlap the test stimulus was examined. Responses to the test stimulus were suppressed by the mask stimulus, which either preceded or followed the test stimulus. To study the temporal characteristics of suppression, the interval between the onset of the test stimulus and that of the mask stimulus (stimulus onset asynchrony, SOA) was varied. Maximum suppression occurred with a simultaneous presentation of the two stimuli, and the suppression gradually weakened as the SOA increased. The suppressive effect of the mask stimulus lasted on average about 77 ms in the negative SOA (forward masking) and 65 ms in the positive SOA (backward masking). These results indicate that surround suppression in V4 neurons has considerable temporal width, which is longer than that previously reported in areas V1 and V2. There were marked differences between the time course of suppression in V4 neurons in the present study and those reported in human metacontrast masking.     

Tracing sequential waves of rapid visuomotor activation in lateralized readiness potentials

Feedforward activation processes are widely regarded as crucial for the automatic initiation of motor responses, whereas recurrent processes are often regarded as crucial for visual awareness. Here, we used a set of behavioral criteria to evaluate whether rapid processing in the human visuomotor system proceeds as would be expected of a feedforward system that works independent of visual awareness. We measured lateralized readiness potentials (LRPs) for key-press responses to color targets ("masks") preceded by masked color primes mapped onto the same or opposite response, and traced the time-course of motor activation as a function of different prime and mask characteristics. LRP time-courses showed that initial motor activation occurred in prime direction and was time-locked to prime onset. Response activation was then captured on-line by the mask signal, with motor activation now time-locked to the mask and proceeding in mask direction. Crucially, the time-course of early activation by the prime was independent of all mask characteristics. This invariance in early priming effects contrasted with large differences in visual awareness for the prime produced by different masks. Results suggest that primed responses to color stimuli are controlled by feedforward waves of activation sequentially elicited by prime and mask signals traveling rapidly enough to escape the recurrent processes leading to backward masking.     

Neuronal correlates of visibility and invisibility in the primate visual system

A brief visual target stimulus may be rendered invisible if it is immediately preceded or followed by another stimulus. This class of illusions, known as visual masking, may allow insights into the neural mechanisms that underlie visual perception. We have therefore explored the temporal characteristics of masking illusions in humans, and compared them with corresponding neuronal responses in the primary visual cortex of awake and anesthetized monkeys. Stimulus parameters that in humans produce forward masking (in which the mask precedes the target) suppress the transient on-response to the target in monkey visual cortex. Those that produce backward masking (in which the mask comes after the target) inhibit the transient after-discharge, the excitatory response that occurs just after the disappearance of the target. These results suggest that, for targets that can be masked (those of short duration), the transient neuronal responses associated with onset and turning off of the target may be important in its visibility.     

The functional relationship between visual backward masking and prepulse inhibition

This experiment demonstrated a relationship between prepulse inhibition (PPI) of the startle eyeblink and visual backward masking in college students. It was hypothesized that recovery from backward masking effects is due in part to sensory gating, as assessed by auditory and visual PPI. Visual presentations of letters served as targets or visual prepulses in an intermixed session of backward masking and PPI. Backward masking and PPI (both auditory and visual) were assessed at stimulus onset asynchronies of 30, 45, 60, 120, and 150 ms. A repeated-measures regression revealed that there was a relationship between backward masking and PPI for both visual and auditory PPI, with higher levels of PPI being associated with greater recovery from backward masking. The data are consistent with the hypothesis that recovery from backward masking effects is affected by sensory gating acting in part to gate out the interruptive effects of the mask.     

Masking selected sequence variation by incorporating mismatches into melting analysis probes

Hybridization probe melting analysis can be complicated by the presence of sequence variation (benign polymorphisms or other mutations) near the targeted mutation. We investigated the use of "masking" probes to differentiate alleles with similar probe melting temperatures. Selected sequence variation was masked by incorporating mismatches (deletion, unmatched nucleotide, or universal base) into hybridization probes at the polymorphic location. Such masking probes create a probe/target mismatch with all possible alleles at the selected polymorphic location. Any allele with additional variation at another site is identified by a lower probe melting temperature than alleles that vary only at the masked position. This "masking technique" was applied to RET protooncogene and HPA6 mutation detection using unlabeled hybridization probes, a saturating dsDNA dye, and high-resolution melting analysis. Masking probes clearly distinguished all targeted mutations from polymorphisms when at least 1 base pair (bp) separated the mutation from the masked variation. We were able to mask polymorphisms immediately adjacent to mutations, except in certain cases, such as those involving single-base deletion probes when both adjacent positions had the same polymorphic nucleotides. The masking probes can also localize mutations to specific codons or nucleotide positions. Masking probes can simplify melting analysis of complex regions and eliminate the need for sequencing.     

Scatter-glare correction using a convolution algorithm with variable weighting

Several investigators have approximated the scatter component of digital radiographic images by applying a constant weighting factor to a convolved version of the detected image. The scatter approximation is then subtracted from the detected image. When a constant weighting factor is used, the scatter in areas of low transmission is underestimated. We have extended this technique by allowing for a spatially variable weighting factor based on the local image intensity. This technique improves the scatter estimate and should provide better results for general videodensitometric applications.     

Spatial localization: tests of a two-process model

The present study examined the recently proposed two-process model of localization performance in which a shift of attention, providing coarse location information, is followed by a saccadic eye movement, providing fine location information. In experiment 1 the nature of the localization response was manipulated. In contrast to the indirect response mode used in the study by Adam et al., i.e., manipulating the arrow keys to move the cursor to the target location, experiment 1 required subjects to point to the target location. The high degree of similarity between the pattern of results obtained with the pointing and cursor response indicated that performance in the localization paradigm was not differentially affected by the nature of the required response. In experiment 2 the characteristics of the backward masking stimulus was manipulated by employing three masking conditions: (1) a long-duration mask; (2) a short-duration mask (100 ms); and (3) a nomask condition. Results showed that the long-duration mask caused interference at short and facilitation at long intervals between onset of target and mask; the short-duration mask caused interference only at short intervals. Overall the findings were consistent with the two-process model of localization performance.     

Pupillary responses on the visual backward masking task reflect general cognitive ability.

Cognitive processing efficiency requires both an ability to attend to task-relevant stimuli with quickness and accuracy, also while filtering distracting or task-irrelevant stimuli. This study investigated cognitive processing efficiency by using pupillary responses as an index of attentional allocation to relevant target and irrelevant masks on a visual backward masking task. The relationship between attentional allocation on this task and general cognitive ability on the scholastic aptitude test (SAT) was examined in college students (n=67). A principle components analysis of the pupillary response waveform isolated a late component that appeared to index the attentional demands associated with processing masks on the backward masking task. This pupillary response index of wasteful resource allocation to the mask accounted for significant variance in SAT scores over and above that accounted for by socio-economic status and target detection accuracy scores. Consistent with the neural efficiency hypothesis, individuals who allocated more resources to processing irrelevant information performed more poorly on cognitive ability tests.     

Foveal versus parafoveal scanpaths of visual imagery in virtual hemianopic subjects

Sophisticated string analysis [compressed regional string analysis, cRSE] shows significant differences following therapeutical masking of the foveal region during a virtual hemianopia. The visual imagery scanpath is done over a compressed mental image that needs longer fixation duration but fewer saccades than the real image. Combination of different viewing tasks with types of pictures permits to show how scanpath top-down strategies can be enforced or decreased by proper combination of task and picture; this is influenced by the mask of fovea versus no-mask of fovea difference, with bottom up mechanisms becoming more important with loss of foveal viewing strategies in the mask condition.     

Application of the EM algorithm to radiographic images.

The expectation maximization (EM) algorithm has received considerable attention in the area of positron emitted tomography (PET) as a restoration and reconstruction technique. In this paper, the restoration capabilities of the EM algorithm when applied to radiographic images is investigated. This application does not involve reconstruction. The performance of the EM algorithm is quantitatively evaluated using a "perceived" signal-to-noise ratio (SNR) as the image quality metric. This perceived SNR is based on statistical decision theory and includes both the observer's visual response function and a noise component internal to the eye-brain system. For a variety of processing parameters, the relative SNR (ratio of the processed SNR to the original SNR) is calculated and used as a metric to compare quantitatively the effects of the EM algorithm with two other image enhancement techniques: global contrast enhancement (windowing) and unsharp mask filtering. The results suggest that the EM algorithm's performance is superior when compared to unsharp mask filtering and global contrast enhancement for radiographic images which contain objects smaller than 4 mm.     

Recognition of lateralized halftone and outline images of everyday objects in conditions of masking

Recognition of the shapes of halftone and outline images of everyday objects in conditions of lateralized tachystoscopic presentation and different levels of noise masking (with “raindrops”) by humans was studied. Mean group data for 15 subjects demonstrated significantly better recognition of outline images of everyday objects by the left hemisphere of the brain than the right at all levels of masking. Increases in masking produced gradual and significant degradation of recognition as compared with controls (recognition of unmasked figures). Recognition of outline images at all levels of masking was significantly better than recognition of halftone images of the same objects. In men, there were no significant differences between hemispheres either at different levels of masking or for different types of stimuli. The neurophysiological mechanisms and functional significance of these effects are discussed. Translated from Zhurnal Vysshei Nervnoi Deyatel’nosti imeni I. P. Pavliva, Vol. 58, No. 1, pp. 56–62, January–February, 2008.     

Velocity selectivity in the cat visual system. II. Independence from interactions between different loci

To investigate the dependence of velocity characteristics on spatiotemporal interactions the velocity selectivity of 15 geniculate and 72 cortical cells (areas 17 and 18) was examined with light and dark bars before and after masking all but the most sensitive part of the receptive field. The use of a 0.3 degree window proved effective in eliminating enough spatiotemporal interactions to abolish cortical direction selectivity. The same window improved the responsiveness at high velocities in only 26% of the cortical cells preferring low velocities and having a receptive field with nonoverlapping ON and OFF subregions. The remaining 74% showed various degrees of velocity-independent decrease in response amplitude. The only two geniculate cells that had a velocity upper cutoff lost this cutoff when tested with the mask. Cortical units preferring high velocities lost their responsiveness at high velocities in the mask condition, provided that their receptive fields contained nonoverlapping ON and OFF subregions. Cortical units, which responded best at intermediate velocities and which had receptive fields with nonoverlapping subregions, lost their sharp velocity tuning when tested with a mask. We conclude that inhibitory spatiotemporal interactions can account for the preference for low velocities in at most a quarter of the cells with nonoverlapping subregions. In contrast, facilitatory interactions seem to be important for cortical cells preferring high or intermediate velocities and having receptive fields with nonoverlapping subregions.     

Pupillometric measures of attentional allocation to target and mask processing on the backward masking task in schizophrenia

Allocation of attentional resources during early visual processing was investigated in schizophrenia. Pupillary responses were recorded during a backward masking task as an index of resource allocation in schizophrenia patients ( n =51) and nonpsychiatric controls ( n =51). Two time-linked components of pupillary response waveforms appeared to differentially index resource allocation to targets versus masks. Two patient subgroups were identified: One with normal overall pupillary responses (resource allocation), but greater allocation on mask relative to target components, and another with abnormally small overall pupillary responses and similar allocation between target and mask components. Thus, misallocation of resources to masks contributed to masking deficits in one subgroup, whereas reduced resource allocation contributed to deficits in the other. The nature of resource-related deficits can vary across schizophrenia subgroups.     

Interactions between IgM antiglobulins and IgG antinuclear antibodies. Some aspects of D-penicillamine activities.

In this report we describe an in vitro masking action of IgM rheumatoid factor (IgM-RF) towards IgG antinuclear antibodies (IgG-ANA) which can be recovered by using D-penicillamine (DP) as an unmasking agent. The mechanism of this masking effect was elucidated by using smears of rat free nucleus as substrate, instead of the classical rat liver cryostat sections technique. It was postulated that the 'inhibition' of IgG-ANA by IgM-RF may be due to the formation of high molecular weight complexes (HMWC); this would be the same in vivo. Allowing the formation of HMWC which can be removed from the circulation, IgM-RF may have a protective effect by preventing or minimizing systemic lesions. Therefore, IgM-RF may be considered as a defence response against potentially noxious ANA or antigen-antibody complexes. Induced nephropathy and the high frequency of detection of ANA observed in rheumatoid arthritis (RA) patients treated by DP may be the result of the dissociating effect of this drug on HMWC, in which the activity of pre-existing ANA is hidden when rat liver sections are used for its detection.     

Transcranial magnetic stimulation in the visual system. II. Characterization of induced phosphenes and scotomas

Transcranial magnetic stimulation (TMS) induces phosphenes and disrupts visual perception when applied over the occipital pole. Both the underlying mechanisms and the brain structures involved are still unclear. In the first part of this study we show that the masking effect of TMS differs to masking by light in terms of the psychometric function. Here we investigate the emergence of phosphenes in relation to perimetric measurements. The coil positions were measured with a stereotactic positioning device, and stimulation sites were characterized in four subjects on the basis of individual retinotopic maps measured by with functional magnetic resonance imaging. Phosphene thresholds were found to lie a factor of 0.59 below the stimulation intensities required to induce visual masking. They covered the segments in the visual field where visual suppression occurred with higher stimulation intensity. Both phosphenes and transient scotomas were found in the lower visual field in the quadrant contralateral to the stimulated hemisphere. They could be evoked from a large area over the occipital pole. Phosphene contours and texture remained quite stable with different coil positions over one hemisphere and did not change with the retinotopy of the different visual areas on which the coil was focused. They cannot be related exclusively to a certain functionally defined visual area. It is most likely that both the optic radiation close to its termination in the dorsal parts of V1 and back-projecting fibers from V2 and V3 back to V1 generate phosphenes and scotomas.     

Effects of perceptual learning in visual backward masking on the responses of macaque inferior temporal neurons

Learning is critical for fast and efficient object recognition in primates. To understand the neuronal correlates of behavioral improvements due to training, we recorded the responses of single neurons in the inferior temporal (IT) cortex of monkeys that were trained to recognize briefly presented, backward-masked objects. First we investigated training effects that are specific to the objects shown during training and that do not transfer to untrained objects. Only one of two monkeys tested showed object-specific training effects at the behavioral level, and only this monkey showed a transient object-specific increase in object selectivity for trained compared with untrained backward-masked objects. However, in each monkey a substantial part of the training effect transferred to untrained objects. To investigate the neural correlates of these object-independent training effects, we compared the neural responses to masked objects in trained monkeys to the responses in untrained monkeys. Training was associated with a reduction of the responses to the irrelevant masking patterns. These findings suggest that extensive training in recognizing backward-masked objects results in neural changes that reduce IT responses to the interfering irrelevant masking patterns and enhance the processing of the relevant objects.     

Contextual influence on orientation discrimination of humans and responses of neurons in V1 of alert monkeys

We studied the effects of various patterns as contextual stimuli on human orientation discrimination, and on responses of neurons in V1 of alert monkeys. When a target line is presented along with various contextual stimuli (masks), human orientation discrimination is impaired. For most V1 neurons, responses elicited by a line in the receptive field (RF) center are suppressed by these contextual patterns. Orientation discrimination thresholds of human observers are elevated slightly when the target line is surrounded by orthogonal lines. For randomly oriented lines, thresholds are elevated further and even more so for lines parallel to the target. Correspondingly, responses of most V1 neurons to a line are suppressed. Although contextual lines inhibit the amplitude of orientation tuning functions of most V1 neurons, they do not systematically alter the tuning width. Elevation of human orientation discrimination thresholds decreases with increasing curvature of masking lines, so does the inhibition of V1 neuronal responses. A mask made of straight lines yields the strongest interference with human orientation discrimination and produces the strongest suppression of neuronal responses. Elevation of human orientation discrimination thresholds is highest when a mask covers only the immediate vicinity of the target line. Increasing the masking area results in less interference. On the contrary, suppression of neuronal responses in V1 increases with increasing mask size. Our data imply that contextual interference observed in human orientation discrimination is in part directly related to contextual inhibition of neuronal activity in V1. However, the finding that interference with orientation discrimination is weaker for larger masks suggests a figure-ground segregation process that is not located in V1.     

Orientational selectivity of the human visual system

1. It is known that an object is less detectable when it is viewed against a background containing structures similar to the object. The effect of changing the orientation between the object and background is investigated.

2. Gratings of variable contrast were generated on two oscilloscopes; these were superimposed optically. The angle of orientation between them could be changed. The threshold of one grating, the test grating, was determined in the presence of the other, the masking grating.

3. When the gratings were presented with the same orientation (and locked in phase) the increment threshold of the test grating was found to be proportional to the suprathreshold contrast of the masking grating.

4. As the angle between the test and masking gratings was increased the masking effect fell exponentially.

5. At 12° on either side of a vertical test grating the masking effect was reduced by a factor of two with respect to its maximum value. This angle was independent of the contrast level of masking, the focus, and also the phase coherence of the masking grating.

6. If the test grating was presented obliquely the effect of masking was slightly less.

7. The narrow orientationally tuned channels found psychophysically by this masking technique are compared with the orientationally sensitive cells discovered electrophysiologically in the visual cortex of the cat.