A model of visual backward masking

When two successive stimuli are presented within 0-200 ms intervals, the recognition of the first stimulus (the target) can be impaired by the second (the mask). This backward masking phenomenon has a form called metacontrast masking where the target and the mask are in close spatial proximity but not overlapping. In that case, the masking effect is strongest for interval of 60-100 ms. To understand this behaviour, activity propagation in a feedforward network of leaky integrate and fire neurons is investigated. It is found that, if neurons have a selectivity similar to that of V1 simple cells, activity decays layer after layer and ceases to propagate. To combat this, a local amplification mechanism is included in the model, using excitatory lateral connections, which turn out to support prolonged self-sustained activity. Masking is assumed to arise from local competition between representations recruited by the target and the mask. This tends to interrupt sustained firing, while prolonged retinal input tends to re-initiate it. Thus, masking causes a maximal reduction of the duration of the cortical response to the target towards the end of the retinal response. This duration exhibits the typical U-shape of the masking curve. In this model, masking does not alter the propagation of the onset of the response to the target, thus preserving response reaction times and enabling unconscious priming phenomena.     

Modulation of activity in human visual area V1 during memory masking

Neurons in the primary visual cortex, V1, are specialized for the processing of elemental features of the visual stimulus, such as orientation and spatial frequency. Recent fMRI evidence suggest that V1 neurons are also recruited in visual perceptual memory; a number of studies using multi-voxel pattern analysis have successfully decoded stimulus-specific information from V1 activity patterns during the delay phase in memory tasks. However, consistent fMRI signal modulations reflecting the memory process have not yet been demonstrated. Here, we report evidence, from three subjects, that the low V1 BOLD activity during retention of low-level visual features is caused by competing interactions between neural populations coding for different values along the spectrum of the dimension remembered. We applied a memory masking paradigm in which the memory representation of a masker stimulus interferes with a delayed spatial frequency discrimination task when its frequency differs from the discriminanda with ±1 octave and found that impaired behavioral performance due to masking is reflected in weaker V1 BOLD signals. This cross-channel inhibition in V1 only occurs with retinotopic overlap between the masker and the sample stimulus of the discrimination task. The results suggest that memory for spatial frequency is a local process in the retinotopically organized visual cortex.     

Forward and backward masking in motor systems

Masking in motor systems was defined as the omission of one act in a sequence due to an earlier or later act in the sequence. A study of phoneme omission in natural speech showed that:

1. 1.

   Masked phonemes were usually preceded or followed by an identical phoneme (referred to as the masking phoneme).

2. 2.

   Backward masking, where the masked phoneme preceded the masking phoneme was as frequent as forward masking.

3. 3.

   The phonemes immediately adjacent to the masked and masking phonemes were usually similar in distinctive features, but rarely identical.

4. 4.

   The masking phoneme usually occurred in a stressed syllable and the masked phoneme in an unstressed one, suggesting that motor intensity may be a factor in masking.

5. 5.

   The components for an adequate model of motor masking were shown to be similar to those in models of other types of errors in speech.

     

Use of backward masking test for the study of visual information processing in healthy subjects and schizophrenic patients

The study of 14 healthy subjects and 15 schizophrenic patients was conducted under visual backward masking conditions. Sensory thresholds were identified using the method of constant stimuli. A special modification of the backward masking technique with lateralized presentation of test and masking stimuli was used to study the lateral characteristics of visual attention. It was found that the thresholds of letter stimulus identification were significantly higher in patients with schizophrenia than in healthy subjects. Only in patients the asymmetry of visual perception was revealed with the higher recognition thresholds in the left visual hemifield. The overall data analysis suggests an association between increased recognition thresholds in schizophrenic patients and changes in the interruption mechanism functioning at the neocortex level.     

A neural network model for selective attention in visual pattern recognition

A neural network model of the mechanism of selective attention in visual pattern recognition is proposed and simulated on a digital computer.When a complex figure consisting of two patterns or more is presented to the model, it is segmented into individual patterns, and each pattern is recognized separately. Even if one of the patterns to which the model is paying selective attention is affected by noise or defects, the model can recall the complete pattern from which the noise has been eliminated and the defects corrected. It is not necessary for perfect recall that the stimulus pattern should be identical in shape to the training pattern. Even though the pattern is distorted in shape or changed in size, it can be correctly recognized and the missing portions restored.The model consists of a hierarchical neural network which has efferent as well as afferent connections between cells. The afferent and the efferent signals interact with each other in the network: the efferent signals, that is, the signals for selective attention, have a facilitating effect on the afferent ones, and, at the same time, the afferent signals gate efferent signal flow. When some feature in the stimulus is not extracted in the afferent paths, the threshold for detection of that feature is automatically lowered by decreasing the efficiency of inhibition, and the model tries to extract even vague traces of the undetected feature.     

Modulation of visual processing by attention and emotion: windows on causal interactions between human brain regions

Visual processing is not determined solely by retinal inputs. Attentional modulation can arise when the internal attentional state (current task) of the observer alters visual processing of the same stimuli. This can influence visual cortex, boosting neural responses to an attended stimulus. Emotional modulation can also arise, when affective properties (emotional significance) of stimuli, rather than their strictly visual properties, influence processing. This too can boost responses in visual cortex, as for fear-associated stimuli. Both attentional and emotional modulation of visual processing may reflect distant influences upon visual cortex, exerted by brain structures outside the visual system per se. Hence, these modulations may provide windows onto causal interactions between distant but interconnected brain regions. We review recent evidence, noting both similarities and differences between attentional and emotional modulation. Both can affect visual cortex, but can reflect influences from different regions, such as fronto-parietal circuits versus the amygdala. Recent work on this has developed new approaches for studying causal influences between human brain regions that may be useful in other cognitive domains. The new methods include application of functional magnetic resonance imaging (fMRI) and electroencephalography (EEG) measures in brain-damaged patients to study distant functional impacts of their focal lesions, and use of transcranial magnetic stimulation concurrently with fMRI or EEG in the normal brain. Cognitive neuroscience is now moving beyond considering the putative functions of particular brain regions, as if each operated in isolation, to consider, instead, how distinct brain regions (such as visual cortex, parietal or frontal regions, or amygdala) may mutually influence each other in a causal manner.     

Modulation of semantic processing by spatial selective attention

The effects of spatial selective attention upon ERPs associated with the processing of word stimuli were investigated. While subjects maintained central eye fixation, ERPs were recorded to words presented to the left and right visual fields. In each of 6 runs, subjects focussed attention to alternate fields to perform a category-detection task. Pairs of semantically related and repeated words were embedded in the word lists presented to the attended and unattended visual fields. Consistent with prior studies, the P1-N1 visual ERP was larger when elicited by words in attended spatial locations. A large negative slow wave identified as N400 was elicited by attended, but not unattended, words. For attended words, N400 was smaller for semantically primed or repeated words. We concluded that spatial selective attention can modulate the degree to which words are processed, and that the cognitive processes associated with N400 are not automatic.     

Analysis and test of laws for backward (metacontrast) masking

In backward visual masking, it is common to find that the mask has its biggest effect when it follows the target by several tens of milliseconds. Research in the 1960s and 1970s suggested that masking effects were best characterized by the stimulus onset asynchrony (SOA) between the target and mask. In particular, one claim has been that the SOA for which masking is optimal remains fixed, even as target and mask durations varied. Experimental evidence supported this claim, and it was accepted as an SOA law. However, recent modeling (Francis, 1997) and experimental studies (Macknik and Livingstone, 1998) argued for new ISI (interstimulus interval) and STA (stimulus termination asynchrony) laws, respectively. This paper reports a mathematical analysis and experimental tests of the laws. The mathematical analysis demonstrates unsuspected relationships between the laws. The experiments test the predictions of the SOA, ISI, and STA laws. The data favor the ISI law over the SOA and the STA laws.     

Computational modelling of visual attention

Five important trends have emerged from recent work on computational models of focal visual attention that emphasize the bottom-up, image-based control of attentional deployment. First, the perceptual saliency of stimuli critically depends on the surrounding context. Second, a unique 'saliency map' that topographically encodes for stimulus conspicuity over the visual scene has proved to be an efficient and plausible bottom-up control strategy. Third, inhibition of return, the process by which the currently attended location is prevented from being attended again, is a crucial element of attentional deployment. Fourth, attention and eye movements tightly interplay, posing computational challenges with respect to the coordinate system used to control attention. And last, scene understanding and object recognition strongly constrain the selection of attended locations. Insights from these five key areas provide a framework for a computational and neurobiological understanding of visual attention.     

Stimulus detection after interruption of the feedforward response in a backward masking paradigm

In backward masking, a target stimulus is rendered invisible by the presentation of a second stimulus, the mask. When the mask is effective, neural responses to the target are suppressed. Nevertheless, weak target responses sometimes may produce a behavioural response. It remains unclear whether the reduced target response is a purely feedforward response or that it includes recurrent activity. Using a feedforward neural network of biological plausible spiking neurons, we tested whether a transient spike burst is sufficient for face categorization. After training the network, the system achieved face/non-face categorization for sets of grayscale images. In a backward masking paradigm, the transient burst response was cut off thereby reducing the feedforward target response. Despite the suppressed feedforward responses stimulus classification remained robust. Thus according to our model data stimulus detection is possible with purely, suppressed feedforward responses.     

Lateralization of visual perception of letters during reverse masking

Identification by healthy subjects of single letters in conditions of backward masking was studied at lateralized presentation of stimuli. Both at dichoptic and unilateral presentations, the letters are identified better if they are presented in the left visual field, i.e. addressed "directly" to the right hemisphere. It is suggested that the data obtained result from the specificity of hemispheres' cooperation under difficult conditions of visual-spatial stage of information processing.     

Effects of set-size and lateral masking in visual search

In the present research, the roles of lateral masking and central processing limitations in visual search were studied. Two search conditions were used: (1) target differed from distractors by presence/absence of a simple feature; (2) target differed by relative position of the same components only. The number of displayed stimuli (set-size) and the distance between neighbouring stimuli were varied as independently as possible in order to measure the effect of both. The effect of distance between stimuli (lateral masking) was found to be similar in both conditions. The effect of set-size was much larger for relative position stimuli. The results support the view that perception of relative position of stimulus components is limited mainly by the capacity of central processing.     

Binocular rivalry requires visual attention

An interocular conflict arises when different images are presented to each eye at the same spatial location. The visual system resolves this conflict through binocular rivalry: observers consciously perceive spontaneous alternations between the two images. Visual attention is generally important for resolving competition between neural representations. However, given the seemingly spontaneous and automatic nature of binocular rivalry, the role of attention in resolving interocular competition remains unclear. Here we test whether visual attention is necessary to?produce rivalry. Using an EEG frequency-tagging method to track cortical representations of the conflicting images, we show that when attention was diverted away, rivalry stopped. The EEG data further suggested that the neural representations of the dichoptic images combined without attention. Thus, attention is necessary for dichoptic images to be engaged in sustained rivalry and may be generally required for resolving conflicting, potentially ambiguous input and giving a single interpretation access to consciousness.     

Analysis of the effect of backward masking by means of a complex model of a net of neuron-like elements

According to the experiments with a projective-associative model of the neuronal net, the phenomenon of “backward masking” of the first stimulus of a pair of stimuli at a small time gap between the stimuli is caused by two events: (1) pre-excitation inhibition of the first stimulus-induced activation by the second stimulus and (2) disturbance of information processing connected with the deficiency of time needed to match the recalled symbol in memory to the symbol presented to the input subsystem and also to name it. Identification of the second stimulus may be impaired with a decreasing time interval due to: (1) superposition of the second (2) recurrent inhibition occurring in the neuronal net upon recognition of the first stimulus. It was found that in conditions of activity of neuron-like elements of the neuronal net, simulating the states of somnolence or slow-wave sleep, corresponding subsystems failed to learn, while time needed to identify already “learned” symbols substantially increased. The data obtained are in agreement with the hypothesis concerning the causes of backward masking and also with the facts on optimal conditions of learning and reproducing its results in living nervous system. It seems reasonable that discussed disturbances of information processing should be kept in mind in designing computers of a new generation, based on the use of principles of brain functioning, in order to increase the reliability and operation speed of technical systems.     

Splitting the spotlight of visual attention

Can the brain attend to more than a single location at one time? In this issue of Neuron, McMains and Somers report psychophysical and fMRI evidence showing that subjects can attend to two separate locations concurrently and that divided spatial attention leads to separate zones of attentional enhancement in early visual cortex.     

Sustained focal attention and peripheral letter recognition

Topographic characteristics of peripheral letter recognition were investigated using a sustained attention paradigm to clarify whether its deployment in the visual field is equally easy in all eight tested locations at 7.5 deg eccentricity. Target size (36 arcmin) was clearly above threshold, so that letters were easily recognized at long durations (> 500 ms). In the main experiment, they were displayed for an individually determined duration of 66 to 167 ms. Six of twelve normally sighted subjects were in their twenties, the others in their fifties. The target location was cued (1 s), and after 2.5-4 s delay, a target was displayed. The results provide strong evidence that performance depended significantly on location and subject. All spatial characteristics showed anisometry, and most showed vertical asymmetry of either sign. Performance was always best on the horizontal meridian. None of the results correlated with subject age. These findings also show that in disfavored locations, performance is limited by deploying attention there, not by holding it there. Consequently, in low vision rehabilitation after binocular central field loss, the choice of a preferred retinal locus for 'eccentric viewing' can be limited by an attentional factor that is unrelated to the eye disease.     

The dark side of visual attention

The limited capacity of neural processing restricts the number of objects and locations that can be attended to. Selected events are readily enhanced: the bright side of attention. However, such focal processing comes at a cost, namely, functional blindness for unattended events: the dark side of visual attention. Recent work has advanced our understanding of the neural mechanisms that facilitate visual processing, as well as the neural correlates of unattended, unconscious visual events. Also, new results have revealed how attentional deployment is optimized by non-visual factors such as behavioral set, past experience, and emotional salience.