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Yet, a subsequent study showed that when manipulating carefully attentional amplification, low-contrast stimuli could induce the same AE whether crowded or not Bi et al. Taken together, these results suggest that crowding may be seen as a bottleneck impacting visual processes differently depending on their position along the visual pathways.

If the independence between AE amplitude and crowding remains debated, the evidence is clearer for CFS: the very first paper introducing CFS showed that it had a detrimental effect on the amplitude of after-images Tsuchiya and Koch, , and recent findings using fMRI strengthened this idea by showing that CFS decreases the activity from signals as early as in the primary visual cortex Watanabe et al. Comparing these results with those of Fang and He described above, one could deduce that crowding impacts visual processes at later stages compared to CFS.

However, considering the high variability across experiments, we argue that clear conclusions should be drawn from systematic comparisons only see discussion. Non-Conscious Processing of Motion Signals As we mentioned in the introduction, CFS and crowding present the advantage of maintaining stimuli out of awareness for potentially long durations.

So far, most studies that took advantage of this property focused on motion processing, by measuring motion AE from invisible stimuli.

In CFS, translational motion has first been shown to induce motion AE that did not transfer across eyes, suggesting that the underlying processes are quite low-level Maruya et al. Yet, with a slightly different setup, AE induced by translational and spiral motion were found to transfer across eyes, and to depend on attentional amplification Kaunitz et al.

Recently, AE from apparent and biological motions i. However, it was found that the extent of temporal integration was smaller under CFS than in condition of full visibility i. Taken together, these results suggest that motion processing is enabled under CFS at various levels of complexity, but with restricted periods of temporal integration.

Interestingly, it was also shown that the processing of implied motion i.

Under crowding, translational motion is known to shift the apparent location of a subsequent stimulus Whitney, ; Bressler and Whitney, , while spiral motion Aghdaee, ; Aghdaee and Zandvakili, and apparent motion Rajimehr et al. In addition, crowded biological motion in the form of dynamic facial expressions were found to be processed despite crowding, notably through the dorsal visual pathway Faivre et al.

It appears from these findings that, as in CFS, motion processing at various levels of complexity is enabled under crowding. Non-Conscious Processing of Multi-Feature Objects Face stimuli Among all stimuli composed of multiple features, faces have undoubtedly triggered the most interest in the field of non-conscious vision, including studies relying on CFS and crowding. Beyond its obvious ecological value, a single face stimulus conveys multiple levels of information which allows for probing non-conscious processing at several levels of complexity.

Here, we review the evidence for non-conscious face processing along two axes: the representation of facial identity and the emotional processing of facial expression. They found that the fusiform face area was more activated by fearful or neutral faces compared to scramble faces.

In addition, they found that the amygdala and superior temporal sulcus were more activated by fearful compared to neutral faces. Recently, it was found that this activation in the superior temporal sulcus was only present in participants with high negative affectivity a dispositional trait relevant to psychopathology, see Vizueta et al.

A subsequent study reported that category-specific responses induced by invisible faces vs. Building upon these fMRI studies, the same research teams then focused on the electromagnetic correlates of non-conscious face processing.

Jiang et al. Sterzer et al. Taken together, these results based on hemodynamic and electromagnetic correlates of neural activity suggest that the signals conveying both face-specific information i. At the behavioral level, the evidence for processing of facial identity under CFS is less convincing.

Using a method similar to CFS, Moradi et al. They found that such identity AE occurred when the adapting face was visible, but completely vanished when it was invisible.

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Later, Stein and Sterzer reported that identity AE could actually be induced by invisible faces, though with a reduced amplitude, and without interocular transfer i. Similarly, Barbot and Kouider found that the identity of invisible faces induced repetition priming effects, but with no interocular transfer: participants were faster to categorize a target face as famous when it was preceded by an invisible identical vs.

As primes and targets were of different sizes i. This raises the possibility that identity priming for faces reflects low-level overlap rather than the activation of face representations per se. In addition, the existence of identity repetition priming in conditions of complete unawareness was recently challenged by a study reporting that priming effects are indeed induced by faces whose identity is invisible, but critically, only when lower level features like color or location are visible i.

We come back to the issue of partial awareness in the discussion. Overall, the behavioral evidence for the processing of facial identity remains inconclusive. Later, it was shown that invisible adaptors depicting facial expressions of anger, fearfulness and happiness could bias the way a subsequent target face was perceived i. Contrary to what was observed for the processing of identity, these results cannot stem from low-level retinotopic similarities, as adaptor and target stimuli were presented to opposite sides of the visual field and to different eyes.

Yet, facial expression AE were subsequently dismissed on the basis that it probably stemmed from residual visibility and attentional confounds Yang et al. Supporting this idea, another study showed that aftereffects from gender and race information were absent under strict control of awareness Amihai et al. New elements to this debate were recently added by two studies, in which invisible fearful faces were found to change skin-conductance responses, both in the context of fear conditioning Raio et al.

In addition, preference judgments biases were found to be induced by angry — but not happy — faces rendered invisible by CFS Almeida et al. We can conclude from this group of recent studies that unlike facial identity, facial expressions rendered invisible by CFS elicit responses that can be captured at the behavioral level 3. Furthermore, we showed that crowded facial expressions can bias subsequent affective judgments of neutral pictographs happy faces elicited more pleasant judgments than angry faces, see Kouider et al.

Moreover, the preference bias induced by crowding faces was not only induced by static i.

Using fMRI coupled with univariate analysis, it was found that compared to a neutral face, static happy faces activated primarily the ventral visual pathway including the fusiform face area, which was functionally connected to the amygdala Faivre et al. By contrast, dynamic happy faces triggered the dorsal visual pathway including the posterior parietal cortex and the substantia innominata, a structure contiguous with the dorsal amygdala. To our knowledge, no multivariate pattern analysis has been applied to try to decode the content of crowded stimuli.

Along the same lines, it was shown that crowded emotional faces could influence a conscious judgment assessing the average emotion resulting of six flanker faces and one target crowded face while the same inverted and scrambled faces could not Haberman and Whitney, , Like our results, this finding shows that despite preventing object recognition, crowding does not impede the processing of emotional information extracted from objects.

Tool stimuli and the dorsal visual stream Besides faces, the processing of tools under CFS has also received much attention. Using fMRI, Fang and He first revealed that suppressed pictures of tools specifically activate the dorsal visual pathway, which is thought to support the guidance of actions Goodale and Milner, At the behavioral level, it was shown that suppressed pictures of tools — but not of non-manipulable objects like animals — could facilitate the categorization of subsequent targets i.

Recently, this finding was challenged by a study revealing that similar priming effects could actually be induced by any kind of elongated objects, rather than specifically manipulable objects Hebart and Hesselmann, ; Sakuraba et al.

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As the previous priming effects may stem from such low-level confounds, the level of processing undergone by invisible tool stimuli remains unclear. Relying on multivariate analysis of blood-oxygen-level dependent signal, Hesselmann and Malach showed that features from invisible tools were encoded in the lateral occipital cortex, which rules out the possibility that stimulus energy was so low that all high-level processes were abolished. Most importantly, they also found that CFS equally reduced brain activity in the ventral and dorsal visual pathways, which challenges the original claim that CFS has no or little influence on the dorsal visual pathway Almeida et al.

The claim that CFS disrupts the ventral but not the dorsal visual pathway is further debated, as one study documented the capacity to grasp visually suppressed stimuli Roseboom and Arnold, , but see Ludwig et al. Regarding crowding, although several lines of evidence suggest that multi-feature objects like arrows Faivre and Kouider, b , sequences of geometric shapes Atas et al.

Yet, it was found that crowding decreases to the same extent the spatial resolution of both visually guided reaching and perception, suggesting that it impacts both the ventral and the dorsal visual pathways Bulakowski et al.

Non-Conscious Processing at the Semantic Level As of today, it remains unclear whether stimuli rendered invisible by CFS can be processed up to the semantic level. Combining the semantic-priming procedure with binocular rivalry, Zimba and Blake first presented prime words to an eye during either dominance or suppression phases of binocular rivalry.

A semantic-priming effect here on response times in a lexical decision task was observed only when prime words were presented during dominance phases, suggesting that semantic processing is disabled during suppression phases. Kang et al. Here, an invisible target word e.

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Although target words usually elicit N components of smaller amplitudes when preceded by semantically related vs. As in binocular rivalry, this result lead the authors to conclude that semantic processing of words was disabled under CFS but see Heyman and Moors, , for possible theoretical and methodological issues.

Contradicting this negative result, it was recently shown that complex — rather than single — semantic stimuli e.

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For instance, the result of an invisible equation e. This suggests that the equation had been non-consciously solved by the time the target appeared. In line with this study, Zabelina et al.

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In each trial, participants had to solve a compound remote associate problem, that is finding a word e. Participants solved word problems faster following suppressed problem words than following suppressed irrelevant words.

Interestingly, this priming effect was observed only when participants reached the solution by analysis rather than by insight, which led the authors to suggest that semantic processing but not semantic integration of the word triads occurred non-consciously.

Here, however, since there was no physical difference between primes and targets i. Indeed, participants may have processed the triplet words at a perceptual but not semantic level, which nevertheless would facilitate responses on the triplet words when they become visible.

In the same vein, using a setting in which participants heard a verbal label before performing a simple detection task wherein stimuli were pictures of familiar objects rendered invisible by CFS, Lupyan and Ward found that valid labels words semantically related to the object improved performance while invalid labels decreased performance. Yet, they also reported that the effectiveness of labels varied as a function of the match between the shape of the stimulus and the shape denoted by the label, suggesting that labels facilitated the perceptual processing of the suppressed objects rather than their semantic processing 4.

Finally, two studies have investigated whether crowded stimuli can be processed at the semantic level. Yeh et al. Recently, Peng et al. The efficacy of our stress induction was established by observing significant increases in cortisol for the stress condition only. We examined extinction retrieval by comparing conditioned responses during the last trial of extinction day 1 with that of the first trial of re-extinction day 2.

Our results suggest that acute stress impairs extinction memory retrieval and offers insight into why treatment strategies used in the clinic may be challenging to recruit in daily life where stress is pervasive.

Keywords: extinction retrieval, fear conditioning, stress, cortisol 1. Introduction The capacity to regulate fear responses to cues as they change from predicting danger to safety is critical to shaping healthy, adaptive behavior.

Extinction learning refers to the process by which conditioned fear responses to a cue conditioned stimulus, or CS that has acquired affective properties after being paired with an aversive outcome unconditioned stimulus, or US gradually diminish as the cue is repeatedly presented without reinforcement Pavlov, ; Bouton, Research across species has shown that the failure to retrieve extinction training can occur over time spontaneous recovery , after a change in context renewal , or after exposure to the US reinstatement Bouton, Another important, but relatively unexplored, factor that may promote impaired extinction retrieval is exposure to stress.

Importantly, the brain regions that support extinction learning and retrieval overlap extensively with those that modulate neuroendocrine responses to stress, rendering these processes especially sensitive to the influence of stress. Nonetheless, the relationship between acute stress and the expression of extinction memory has not been directly examined in humans. Stress and stress hormones i.

For example, across a wide range of declarative and episodic learning and memory tasks, stress and stress hormones i. Neurobiological research in non-human animals has provided a detailed understanding of how stress interacts with the neural circuitry that supports each phase of cued-based fear conditioning and extinction, offering insight into how stress can differentially affect the way in which aversive associations are learned, extinguished and later retrieved.

The acquisition, consolidation and retrieval of cued fear conditioning rely on the amygdala, which serves as the site of CS-US convergence LeDoux ; Maren Consistent with this, work in rodents has shown stress exposure or administration of glucocorticoids facilitates both the acquisition Wilson et al, ; Shors et al, ; Shors, and consolidation Hui et al.

The consolidation of extinction learning, however, is enhanced with glucocorticoid administration, suggesting that despite the variable effects of stress on extinction learning, stress hormones facilitate the long-term storage of such training Cai et al ; Yang et al, Chronic stress Miracle et al, ; Garcia et al, ; Wilber et al, and single, prolonged stress exposure Knox et al, has been shown to impair extinction retention after intact training.

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