Recent models and findings in visual backward masking: a comparison, review, and update

Visual backward masking not only is an empirically rich and theoretically interesting phenomenon but also has found increasing application as a powerful methodological tool in studies of visual information processing and as a useful instrument for investigating visual function in a variety of specific subject populations. Since the dual-channel, sustained-transient approach to visual masking was introduced about two decades ago, several new models of backward masking and metacontrast have been proposed as alternative approaches to visual masking. In this article, we outline, review, and evaluate three such approaches: an extension of the dual-channel approach as realized in the neural network model of retino-cortical dynamics (Ogmen, 1993), the perceptual retouch theory (Bachmann, 1984, 1994), and the boundary contour system (Francis, 1997; Grossberg & Mingolla, 1985b). Recent psychophysical and electrophysiological findings relevant to backward masking are reviewed and, whenever possible, are related to the aforementioned models. Besides noting the positive aspects of these models, we also list their problems and suggest changes that may improve them and experiments that can empirically test them.     

A comparison of masking by visual and transcranial magnetic stimulation: implications for the study of conscious and unconscious visual processing

Visual stimuli as well as transcranial magnetic stimulation (TMS) can be used: (1) to suppress the visibility of a target and (2) to recover the visibility of a target that has been suppressed by another mask. Both types of stimulation thus provide useful methods for studying the microgenesis of object perception. We first review evidence of similarities between the processes by which a TMS mask and a visual mask can either suppress the visibility of targets or recover such suppressed visibility. However, we then also point out a significant difference that has important implications for the study of the time course of unconscious and conscious visual information processing and for theoretical accounts of the processes involved. We present evidence and arguments showing: (a) that visual masking techniques, by revealing more detailed aspects of target masking and target recovery, support a theoretical approach to visual masking and visual perception that must take into account activities in two separate neural channels or processing streams and, as a corollary, (b) that at the current stage of methodological sophistication visual masks, by acting in more highly specifiable ways on these pathways, provide information about the microgenesis of form perception not available with TMS masks.     

Unconscious priming by color and form: different processes and levels

Using a metacontrast masking paradigm, prior studies have shown (a) that a target's color information and form information, can be processed without awareness and (b) that unconscious color processing occurs at early, wavelength-dependent levels in the cortical information processing hierarchy. Here we used a combination of paracontrast and metacontrast masking techniques to explore unconscious color and form priming effects produced by blue, green, and neutral stimuli. We found that color priming in normal observers is significantly reduced when an additional paracontrast mask precedes the target at optimal masking SOAs. However, no reduction of form-priming effects was obtained at similar optimal paracontrast SOAs. We conclude that unconscious color priming depends on an early, wavelength- or stimulus-dependent response of color neurons located at early cortical levels whereas unconscious form priming occurs at later levels.     

Unconscious, stimulus-dependent priming and conscious, percept-dependent priming with chromatic stimuli

Using metacontrast masking to suppress the conscious registration of a prime stimulus, Breitmeyer, Ro, and Singhal (2004) showed that color priming produced by a masked prime disk occurs at unconscious stimulus-dependent rather than at percept-dependent levels of visual processing. The current set of experiments compares this type of unconscious stimulus-dependent priming to conscious priming produced by a prime that, in two separate ways, is rendered visible and thus activates percept-dependent visual processes. The results indicate that while the masked prime again acts at a stimulus-dependent level of processing, the unmasked, visible primes additionally act at a later percept-dependent level of processing.     

Exogenous attention during perceptual group formation and dissolution

Under natural viewing conditions, a large amount of information reaches our senses, and the visual system uses attention and perceptual grouping to reduce the complexity of stimuli in order to make real-time perception possible. Prior studies have shown that attention and perceptual grouping operate in synergy; exogenous attention is deployed not only to the cued item, but also to the entire group. Here, we investigated how attention and perceptual grouping operate during the formation and dissolution of groups. Our results showed that reaction times are higher in the presence of perceptual groups than they are for ungrouped stimuli. On the other hand, attentional benefits of perceptual grouping were observed during both the formation and the dissolution of groups. The dynamics were similar during group formation and dissolution, showing a gradual effect that takes approximately half a second to reach its maximum level. In the case of group dissolution, the attentional benefits persisted for about a quarter of a second after dissolution of the group. Taken together, our results reveal the dynamics of how attention and grouping work in synergy during the transient period when groups form or dissolve.     

Differential effects of inactivation of the orbitofrontal cortex on strategy set-shifting and reversal learning

Different subregions of the rodent prefrontal cortex (PFC) mediate dissociable types of behavioral flexibility. For example, lesions of the medial or orbitofrontal (OFC) regions of the PFC impair extradimensional shifts and reversal learning, respectively, when novel stimuli are used during different phases of the task. In the present study, we assessed the effects of inactivation of the OFC on strategy set-shifting and reversal learning, using a maze based set-shifting task mediated by the medial PFC. Long–Evans rats were trained initially on a visual-cue discrimination to obtain food. On the subsequent day, rats had to shift to using a response strategy (e.g., always turn left). On Day 3 (reversal), rats were required to reverse the direction of their turn (e.g., always turn right). Infusions of the local anesthetic bupivacaine into the OFC did not impair initial visual discrimination learning, nor did it impair performance on the set-shift. In contrast, inactivation of the OFC did impair reversal learning; yet, these rats ceased using the previously acquired response rule as readily as controls. Instead, rats receiving OFC inactivations made a disproportionate number of erroneous arm entries towards the visual-cue, suggested that these animals reverted back to using the original visual-cue based strategy. These findings, in addition to previous data, further support the notion that the OFC and medial PFC play dissociable roles in reversal learning and set-shifting. Furthermore, the lack of effect of OFC inactivations on the set-shift indicates that this type of behavioral flexibility does not require cognitive operations related to reversal learning.     

Effects of contrast polarity in paracontrast masking

The visibility of a target stimulus can be suppressed (inhibition) or increased (facilitation) during paracontrast masking. Three processes have been proposed to be involved in paracontrast masking: brief inhibition, facilitation, and prolonged inhibition (Breitmeyer et al., 2006). Brief inhibition is observed when the mask precedes the target at short stimulus onset asynchronies (SOAs) ranging from -10 to -30 msec, whereas prolonged inhibition is effective up to very large SOAs of -450 msec. Facilitation, enhancement in target visibility, can be observed at SOA values between -20 and -110 msec. We further investigated these processes by changing target-mask spatial separation and the contrast polarity of the mask. Our results show that (1) facilitation weakens when spatial separation between the target and mask is increased or when they have opposite contrast polarity, and (2) brief inhibition turns into facilitation for the opposite-polarity mask, whereas prolonged inhibition does not change significantly. These results suggest a fast inhibition mechanism realized in the contrast-specific center-surround antagonism of classical receptive fields for brief inhibition and a slower, higher level cortical processing that is indifferent to contrast polarity for prolonged inhibition.     

The reference frame for encoding and retention of motion depends on stimulus set size

The goal of this study was to investigate the reference frames used in perceptual encoding and storage of visual motion information. In our experiments, observers viewed multiple moving objects and reported the direction of motion of a randomly selected item. Using a vector-decomposition technique, we computed performance during smooth pursuit with respect to a spatiotopic (nonretinotopic) and to a retinotopic component and compared them with performance during fixation, which served as the baseline. For the stimulus encoding stage, which precedes memory, we found that the reference frame depends on the stimulus set size. For a single moving target, the spatiotopic reference frame had the most significant contribution with some additional contribution from the retinotopic reference frame. When the number of items increased (Set Sizes 3 to 7), the spatiotopic reference frame was able to account for the performance. Finally, when the number of items became larger than 7, the distinction between reference frames vanished. We interpret this finding as a switch to a more abstract nonmetric encoding of motion direction. We found that the retinotopic reference frame was not used in memory. Taken together with other studies, our results suggest that, whereas a retinotopic reference frame may be employed for controlling eye movements, perception and memory use primarily nonretinotopic reference frames. Furthermore, the use of nonretinotopic reference frames appears to be capacity limited. In the case of complex stimuli, the visual system may use perceptual grouping in order to simplify the complexity of stimuli or resort to a nonmetric abstract coding of motion information.     

Facilitation by exogenous attention for static and dynamic gestalt groups

Attentional mechanisms allow the brain to selectively allocate its resources to stimuli of interest within the huge amount of information reaching its sensory systems. The voluntary component of attention, endogenous attention, can be allocated in a flexible manner depending on the goals and strategies of the observer. On the other hand, the reflexive component, exogenous attention, is driven by the stimulus. Here, we investigated how exogenous attention is deployed to moving stimuli that form distinct perceptual groups. We showed that exogenous attention is deployed according to a reference frame that moves along with the stimulus. Moreover, in addition to the cued stimulus, exogenous attention is deployed to all elements forming a perceptual group. These properties provide a basis for the efficient deployment of exogenous attention under ecological viewing conditions.