The priming of stimulus-elicited involuntary mental imagery.

Percepts, urges, and even high-level cognitions often enter the conscious field involuntarily. The Reflexive Imagery Task (RIT) was designed to investigate experimentally the nature of such entry into consciousness. In the most basic version of the task, participants are instructed not to subvocalize the names of visual objects. Involuntary subvocalizations arise on the majority of the trials. Can these effects be influenced by priming? In our experiment, participants were exposed to an auditory prime 300 ms before being presented with the RIT stimuli. For example, participants heard the word "FOOD" before seeing two RIT stimuli (e.g., line drawings of BANANA and CAT, with the former being the target of the prime). The short span between prime and target allowed us to assess whether the RIT effect is strategic or automatic. Before each trial, participants were instructed to disregard what they hear, and not to think of the name of any of the objects. On an average of 83% of the trials, the participants thought (involuntarily) of the name of the object associated with the prime. This is the first study to use a priming technique within the context of the RIT. The theoretical implications of these involuntary effects are discussed.

Prime-induced illusion of control: The influence of unconscious priming on self-initiated actions and the role of regression to the mean.

To what degree human cognition is influenced by subliminal stimuli is a controversial empirical question. One striking example was reported by Linser and Goschke (2007): participants overestimated how much control they had over objectively uncontrollable stimuli when masked congruent primes were presented immediately before the action. Critically, however, unawareness of the masked primes was established by post hoc data selection. In our preregistered study we sought to explore these findings while adjusting prime visibility based on individual thresholds, so that each participant underwent both visible and non-visible conditions. In experiment 1, N = 39 participants engaged in a control judgement task: following the presentation of a semantic prime, they freely selected between two keys, which triggered the appearance of a colored circle. The color of the circles, however, was independent of the key-press. Subsequently, participants assessed their perceived control over the circle's color, based on their key-presses, via a rating scale that ranged from 0 % (no control) to 100 % (complete control). Contrary to Linser and Goschke (2007)'s findings, this experiment demonstrated that predictive information influenced the experience of agency only when primes were consciously processed. In experiment 2, utilizing symbolic (arrow) primes, N = 35 participants had to rate their feeling of control over the effect-stimulus' identity during a two-choice identification paradigm (i.e., they were instructed to press a key corresponding to a target stimulus; with a contingency between target and effect stimulus of 75 %/25 %). The results revealed no significant influence of subliminal priming on agency perceptions. In summary, this study implies that unconscious stimuli may not exert a substantial influence on the conscious experience of agency, underscoring the need for careful consideration of methodological aspects and experimental design's impact on observed phenomena.

Induced oscillatory brain responses under virtual reality conditions in the context of repetition priming.

In the human electroencephalogram (EEG), induced oscillatory responses in various frequency bands are regarded as valuable indices to examine the neural mechanisms underlying human memory. While the advent of virtual reality (VR) drives the investigation of mnemonic processing under more lifelike settings, the joint application of VR and EEG methods is still in its infancy (e.g., due to technical limitations impeding the signal acquisition). The objective of the present EEG study was twofold. First, we examined whether the investigation of induced oscillations under VR conditions yields equivalent results compared to standard paradigms. Second, we aimed at obtaining further insights into basic memory-related brain mechanisms in VR. To these ends, we relied on a standard implicit memory design, namely repetition priming, for which the to-be-expected effects are well-documented for conventional studies. Congruently, we replicated a suppression of the evoked potential after stimulus onset. Regarding the induced responses, we observed a modulation of induced alphaband in response to a repeated stimulus. Importantly, our results revealed a repetition-related suppression of the high-frequency induced gammaband response (>30 Hz), indicating the sharpening of a cortical object representation fostering behavioral priming effects. Noteworthy, the analysis of the induced gammaband responses required a number of measures to minimize the influence of external and internal sources of artefacts (i.e., the electrical shielding of the technical equipment and the control for miniature eye movements). In conclusion, joint VR-EEG studies with a particular focus on induced oscillatory responses offer a promising advanced understanding of mnemonic processing under lifelike conditions.

Separating the role of perceptual and conceptual fluency on masked word priming using event-related potentials.

Masked word repetition increases "old" responses on an episodic recognition test (Jacoby & Whitehouse, 1989). This effect is commonly attributed to perceptual fluency; that is, unconscious perception of the prime speeds reading of the target and this fluency leads to elevated familiarity. Two experiments directly tested the claim that perceptual fluency is responsible for word priming effects. Experiment 1 held prime-target meaning constant and altered the physical characteristics of match primes (e.g., "RIGHT" primes "RIGHT") by including both lowercase (e.g, "right") and mixed case primes (e.g., "rIgHt"). If word priming effects are due to perceptual fluency, then lowering the perceptual overlap between the prime and target should decrease or eliminate word priming effects. Instead, all three conditions showed robust priming effects in the behavioral and ERP (i.e., N400) measures. Experiment 2 equated the prime-target perceptual features and lowered the conceptual overlap by using orthographically similar nonwords as primes (e.g., "JIGHT" primes "RIGHT"). Removing prime-target conceptual overlap eliminated behavioral evidence of priming and N400 ERP differences correlated with priming. The evidence suggests that word priming effects on episodic recognition memory are more likely a product of conceptual fluency than perceptual fluency.

Attentional priming in Go No-Go search tasks.

Go/No-Go responses in visual search yield different estimates of the operation of visual attention than more standard present versus absent tasks. Such minor methodological tweaks have a surprisingly large effect on measures that have, for the last half-century or so, formed the backbone of prominent theories of visual attention. Secondly, priming effects in visual search have a dominating influence on visual search, accounting for effects that have been attributed to top-down guidance in standard theories. Priming effects in visual search have, however, never been investigated for searches involving Go/No-Go present/absent decisions. Here, Go/No-Go tasks were used to assess visual search for an odd-one-out face, defined either by color or facial expression. The Go/No-Go responses for the color-based task were very fast for both present and absent trials and notably, they resulted in negative slopes of RT and set size. Interestingly "Go" responses were even faster for the target absent case. The "Go" responses were, on the other hand, much slower for expression and became higher with increased set-size, particularly for the target-absent response. Priming effects were considerable for the feature search, but for expression, the target absent priming was strong, but did not occur for target present trials, arguing that repetition priming for this search mainly reflects priming of context rather than target features. Overall, the results reinforce the point that Go/No-Go tasks are highly informative for theoretical accounts of visual attention and are shown here to cast a new light on attentional priming.

Cross-Modal Somatosensory Repetition Priming and Speech Processing.

BACKGROUND: Motor speech treatment approaches have been applied in both adults with aphasia and apraxia of speech and children with speech-sound disorders. Identifying links between motor speech intervention techniques and the modes of action (MoA) targeted would improve our understanding of how and why motor speech interventions achieve their effects, along with identifying its effective components. The current study focuses on identifying potential MoAs for a specific motor speech intervention technique. OBJECTIVES: We aim to demonstrate that somatosensory inputs can influence lexical processing, thus providing further evidence that linguistic information stored in the brain and accessed as part of speech perception processes encodes information related to speech production. METHODS: In a cross-modal repetition priming paradigm, we examined whether the processing of external somatosensory priming cues was modulated by both word-level (lexical frequency, low- or high-frequency) and speech sound articulatory features. The study participants were divided into two groups. The first group consisted of twenty-three native English speakers who received somatosensory priming stimulation to their oro-facial structures (either to labial corners or under the jaw). The second group consisted of ten native English speakers who participated in a control study where somatosensory priming stimulation was applied to their right or left forehead as a control condition. RESULTS: The results showed significant somatosensory priming effects for the low-frequency words, where the congruent somatosensory condition yielded significantly shorter reaction times and numerically higher phoneme accuracy scores when compared to the incongruent somatosensory condition. Data from the control study did not reveal any systematic priming effects from forehead stimulation (non-speech related site), other than a general (and expected) tendency for longer reaction times with low-frequency words. CONCLUSIONS: These findings provide further support for the notion that speech production information is represented in the mental lexicon and can be accessed through exogenous Speech-Language Pathologist driven somatosensory inputs related to place of articulation.

Precise timing of ERK phosphorylation/dephosphorylation determines the outcome of trial repetition during long-term memory formation.

Two-trial learning in Aplysia reveals nonlinear interactions between training trials: A single trial has no effect, but two precisely spaced trials induce long-term memory. Extracellularly regulated kinase (ERK) activity is essential for intertrial interactions, but the mechanism remains unresolved. A combination of immunochemical and optogenetic tools reveals unexpected complexity of ERK signaling during the induction of long-term synaptic facilitation by two spaced pulses of serotonin (5-hydroxytryptamine, 5HT). Specifically, dual ERK phosphorylation at its activating TxY motif is accompanied by dephosphorylation at the pT position, leading to a buildup of inactive, singly phosphorylated pY-ERK. Phosphorylation and dephosphorylation occur concurrently but scale differently with varying 5HT concentrations, predicting that mixed two-trial protocols involving both "strong" and "weak" 5HT pulses should be sensitive to the precise order and timing of trials. Indeed, long-term synaptic facilitation is induced only when weak pulses precede strong, not vice versa. This may represent a physiological mechanism to prioritize memory of escalating threats.

Distinct deficits of repetition priming following lateral versus anteromedial frontal cortex damage.

Object repetition commonly leads to long-lasting improvements in identification speed and accuracy, a behavioral facilitation referred to as "repetition priming". Neuroimaging and non-invasive electromagnetic stimulation studies have most often implicated the involvement of left lateral frontal cortex in repetition priming, although convergent evidence from neuropsychological studies is lacking. In the current study, we examine the impact of surgical resection for the treatment of epilepsy on the magnitude of repetition priming at relatively short-term (30-60 min delay) and long-term (3 months) delays in 41 patients with varying seizure foci and resection locations. Overall, patients exhibited significant repetition priming at both short-term and long-term delays. However, patients with frontal resections (largely anterior and medial frontal) differed significantly from those with right anterior temporal resections in showing fully intact short-term priming but absent long-term priming. In a comparison set of 10 recovered aphasic patients, patients with left lateral frontal damage exhibited impaired short-term priming relative to other frontal damage locations, suggesting the differential involvement of lateral and anteromedial frontal regions in mediating repetition priming at short-lag and long-lag timescales, respectively.

Instructing item-specific switch probability: expectations modulate stimulus-action priming.

Both active response execution and passive listening to verbal codes (a form of instruction) in single prime trials lead to item-specific repetition priming effects when stimuli re-occur in single probe trials. This holds for task-specific classification (stimulus-classification, SC priming, e.g., apple-small) and action (stimulus-action, SA priming, e.g., apple-right key press). To address the influence of expectation on item-specific SC and SA associations, we tested if item-specific SC and SA priming effects were modulated by the instructed probability of re-encountering individual SC or SA mappings (25% vs. 75% instructed switch probability). Importantly, the experienced item-specific switch probability was always 50%. In Experiment 1 (N = 78), item-specific SA/SC switch  expectations affected SA, but not SC priming effects exclusively following active response execution. Experiment 2 (N = 40) was designed to emphasize SA priming by only including item-specific SC repetitions. This yielded stronger SA priming for 25% vs. 75% expected switch probability, both following response execution as in Experiment 1 and also following verbally coded SA associations. Together, these results suggest that SA priming effects, that is, the encoding and retrieval of SA associations, is modulated by item-specific switch expectation. Importantly, this expectation effect cannot be explained by item-specific associative learning mechanisms, as stimuli were primed and probed only once and participants experienced item-specific repetitions/switches equally often across stimuli independent of instructed switch probabilities. This corroborates and extends previous results by showing that SA priming effects are modulated by  expectation not only based on experienced item-specific switch probabilities, but also on mere instruction.

Comprehension exposures to words in sentence contexts impact spoken word production.

Comprehension or production of isolated words and production of words embedded in sentence contexts facilitated later production in previous research. The present study examined the extent to which contextualized comprehension exposures would impact later production. Two repetition priming experiments were conducted with Spanish-English bilingual participants. In Experiment 1 (N = 112), all encoding stimuli were presented visually, and in Experiment 2 (N = 112), all encoding stimuli were presented auditorily. After reading/listening or translating isolated words or words embedded in sentences at encoding, pictures corresponding to each target word were named aloud. Repetition priming relative to new items was measured in RT and accuracy. Relative to isolated encoding, sentence encoding reduced RT priming but not accuracy priming. In reading/listening encoding conditions, both isolated and embedded words elicited accuracy priming in picture naming, but only isolated words elicited RT priming. In translation encoding conditions, repetition priming effects in RT (but not accuracy) were stronger for lower-frequency words and with lower proficiency in the picture-naming response language. RT priming was strongest when the translation response at encoding was produced in the same language as final picture naming. In contrast, accuracy priming was strongest when the translation stimulus at encoding was comprehended in the same language as final picture naming. Thus, comprehension at encoding increased the rate of successful retrieval, whereas production at encoding speeded later production. Practice of comprehension may serve to gradually move less well-learned words from receptive to productive vocabulary.

Tracking the time course of sign recognition using ERP repetition priming.

Repetition priming and event-related potentials (ERPs) were used to investigate the time course of sign recognition in deaf users of American Sign Language. Signers performed a go/no-go semantic categorization task to rare probe signs referring to people; critical target items were repeated and unrelated signs. In Experiment 1, ERPs were time-locked either to the onset of the video or to sign onset within the video; in Experiment 2, the same full videos were clipped so that video and sign onset were aligned (removing transitional movements), and ERPs were time-locked to video/sign onset. All analyses revealed an N400 repetition priming effect (less negativity for repeated than unrelated signs) but differed in the timing and/or duration of the N400 effect. Results from Experiment 1 revealed that repetition priming effects began before sign onset within a video, suggesting that signers are sensitive to linguistic information within the transitional movement to sign onset. The timing and duration of the N400 for clipped videos were more parallel to that observed previously for auditorily presented words and was 200 ms shorter than either time-locking analysis from Experiment 1. We conclude that time-locking to full video onset is optimal when early ERP components or sensitivity to transitional movements are of interest and that time-locking to the onset of clipped videos is optimal for priming studies with fluent signers.

Predictive pre-activation of orthographic and lexical-semantic representations facilitates visual word recognition.

To a crucial extent, the efficiency of reading results from the fact that visual word recognition is faster in predictive contexts. Predictive coding models suggest that this facilitation results from pre-activation of predictable stimulus features across multiple representational levels before stimulus onset. Still, it is not sufficiently understood which aspects of the rich set of linguistic representations that are activated during reading-visual, orthographic, phonological, and/or lexical-semantic-contribute to context-dependent facilitation. To investigate in detail which linguistic representations are pre-activated in a predictive context and how they affect subsequent stimulus processing, we combined a well-controlled repetition priming paradigm, including words and pseudowords (i.e., pronounceable nonwords), with behavioral and magnetoencephalography measurements. For statistical analysis, we used linear mixed modeling, which we found had a higher statistical power compared to conventional multivariate pattern decoding analysis. Behavioral data from 49 participants indicate that word predictability (i.e., context present vs. absent) facilitated orthographic and lexical-semantic, but not visual or phonological processes. Magnetoencephalography data from 38 participants show sustained activation of orthographic and lexical-semantic representations in the interval before processing the predicted stimulus, suggesting selective pre-activation at multiple levels of linguistic representation as proposed by predictive coding. However, we found more robust lexical-semantic representations when processing predictable in contrast to unpredictable letter strings, and pre-activation effects mainly resembled brain responses elicited when processing the expected letter string. This finding suggests that pre-activation did not result in "explaining away" predictable stimulus features, but rather in a "sharpening" of brain responses involved in word processing.

The role of selective attention in the positivity offset: Evidence from event related potentials.

Some research suggests that positive and negative valence stimuli may be processed differently. For example, negative material may capture and hold attention more readily than equally arousing positive material. This is called the negativity bias, and it has been observed as both behavioural and electroencephalographic (EEG) effects. Consequently, it has been attributed to both automatic and elaborative processes. However, at the lowest levels of arousal, faster reaction times and stronger EEG responses to positive material have been observed. This is called the positivity offset, and the underlying cognitive mechanism is less understood. To study the role of selective attention in the positivity offset, participants completed a negative affective priming (NAP) task modified to dissociate priming for positive and negative words. The task required participants to indicate the valence of a target word, while simultaneously ignoring a distractor. In experiment 1, a behavioural facilitation effect (faster response time) was observed for positive words, in stark contrast to the original NAP task. These results were congruent with a previously reported general categorization advantage for positive material. In experiment 2, participants performed the task while EEG was recorded. In additional to replicating the behavioural results from experiment 1, positive words elicited a larger Late Positive Potential (LPP) component on ignored repetition relative to control trials. Surprisingly, negative words elicited a larger LPP than positive words on control trials. These results suggest that the positivity offset may reflect a greater sensitivity to priming effects due to a more flexible attentional set.

Enhanced inter-regional coupling of neural responses and repetition suppression provide separate contributions to long-term behavioral priming.

Stimulus identification commonly improves with repetition over long delays ("repetition priming"), whereas neural activity commonly decreases ("repetition suppression"). Multiple models have been proposed to explain this brain-behavior relationship, predicting alterations in functional and/or effective connectivity (Synchrony and Predictive Coding models), in the latency of neural responses (Facilitation model), and in the relative similarity of neural representations (Sharpening model). Here, we test these predictions with fMRI during overt and covert naming of repeated and novel objects. While we find partial support for predictions of the Facilitation and Sharpening models in the left fusiform gyrus and left frontal cortex, the data were most consistent with the Synchrony model, with increased coupling between right temporoparietal and anterior cingulate cortex for repeated objects that correlated with priming magnitude across participants. Increased coupling and repetition suppression varied independently, each explaining unique variance in priming and requiring modifications of all current models.

Tactile to visual number priming in the left intraparietal cortex of sighted Braille readers.

Numbers can be presented in different notations and sensory modalities. It is currently debated to what extent these formats overlap onto a single representation. We asked whether such an overlap exists between symbolic numbers represented in two sensory modalities: Arabic digits and Braille numbers. A unique group of sighted Braille readers underwent extensive Braille reading training and was tested in an fMRI repetition-suppression paradigm with tactile Braille digit primes and visual Arabic digit targets. Our results reveal cross-modal priming: compared to repetition of two different quantities (e.g., Braille "5" and Arabic "2"), repetition of the same quantity presented in two modalities (e.g., Braille "5" and Arabic "5") led to a reduction of activation in several sub-regions of the Intraparietal Sulcus (IPS), a key cortical region for magnitude processing. Thus, in sighted Braille readers, the representations of numbers read by sight and by touch overlap to a degree sufficient to cause repetition suppression. This effect was modulated by the numerical prime-probe distance. Altogether this indicates that the left parietal cortex hosts neural assemblies that are sensitive to numerical information from different notations (number words or Arabic digits) and modalities (tactile and visual).

Morphosemantic activation of opaque Chinese words in sentence comprehension.

Two cross-modal priming experiments were conducted to investigate morphological processing in Chinese spoken word recognition during sentence comprehension. Participants heard sentences that contained opaque prime words and performed lexical decisions on visual targets that were related to second morpheme meanings of opaque words or whole-word meanings. The targets were presented at the auditory onset of the second morphemes or the subsequent syllables after the opaque primes to examine the time course of effects. In a neutral sentence context (Experiment 1), opaque word morpheme meanings produced morphological priming on target word recognition, which preceded lexical priming. When context biased toward whole opaque words (Experiment 2), morphological priming disappeared, while the effect of lexical meanings remained significant and emerged earlier than the effect of lexical meanings in the neutral context. These findings suggest that morphemes play a role in Chinese spoken word recognition, but their effects depend on the prior context during sentence comprehension.

Age-related changes in repetition suppression of neural activity during emotional future simulation.

Despite advances in understanding the consequences of age-related episodic memory decline for future simulation, much remains unknown regarding changes in the neural underpinnings of future thinking with age. We used a repetition suppression paradigm to explore age-related changes in the neural correlates of emotional future simulation. Younger and older adults simulated positive, negative, and neutral future events either 2 or 5 times. Reductions in neural activity for events simulated 5 versus 2 times (i.e., repetition suppression) identify brain regions responsive to the specific emotion of simulated events. Critically, older adults showed greater repetition suppression than younger adults in the temporal pole for negative simulations, and the cuneus for positive simulations. These findings suggest that older adults distance themselves from negative future possibilities by thinking about them in a more semantic way, consistent with the view that older adults down-regulate negative affect and up-regulate positive affect. More broadly this study increases our understanding of the impact of aging on the neural underpinnings of episodic future simulation.

Pupillometry correlates of visual priming, and their dependency on autistic traits.

In paradigms of visual search where the search feature (say color) can change from trial to trials, responses are faster for trials where the search color is repeated than when it changes. This is a clear example of "priming" of attention. Here we test whether the priming effects can be revealed by pupillometry, and also whether they are related to autistic-like personality traits, as measured by the Autism-Spectrum Quotient (AQ). We repeated Maljkovic and Nakayama's (1994) classic priming experiment, asking subjects to identify rapidly the shape of a singleton target defined by color. As expected, reaction times were faster when target color repeated, and the effect accumulated over several trials; but the magnitude of the effect did not correlate with AQ. Reaction times were also faster when target position was repeated, again independent of AQ. Presentation of stimuli caused the pupil to dilate, and the magnitude of dilation was greater for switched than repeated trials. This effect did not accumulate over trials, and did not correlate with the reaction times difference, suggesting that the two indexes measure independent aspects of the priming phenomenon. Importantly, the amplitude of pupil modulation correlated negatively with AQ, and was significant only for those participants with low AQ. The results confirm that pupillometry can track perceptual and attentional processes, and furnish useful information unobtainable from standard psychophysics, including interesting dependencies on personality traits.

Changes in human brain dynamics during behavioral priming and repetition suppression.

Behavioral responses to a perceptual stimulus are typically faster with repeated exposure to the stimulus (behavioral priming). This implicit learning mechanism is critical for survival but impaired in a variety of neurological disorders, including Alzheimer's disease. Many studies of the neural bases for behavioral priming have encountered an interesting paradox: in spite of faster behavioral responses, repeated stimuli usually elicit weaker neural responses (repetition suppression). Several neurophysiological models have been proposed to resolve this paradox, but noninvasive techniques for human studies have had insufficient spatial-temporal precision for testing their predictions. Here, we used the unparalleled precision of electrocorticography (ECoG) to analyze the timing and magnitude of task-related changes in neural activation and propagation while patients named novel vs repeated visual objects. Stimulus repetition was associated with faster verbal responses and decreased neural activation (repetition suppression) in ventral occipito-temporal cortex (VOTC) and left prefrontal cortex (LPFC). Interestingly, we also observed increased neural activation (repetition enhancement) in LPFC and other recording sites. Moreover, with analysis of high gamma propagation we observed increased top-down propagation from LPFC into VOTC, preceding repetition suppression. The latter results indicate that repetition suppression and behavioral priming are associated with strengthening of top-down network influences on perceptual processing, consistent with predictive coding models of repetition suppression, and they support a central role for changes in large-scale cortical dynamics in achieving more efficient and rapid behavioral responses.

Verbing nouns and nouning verbs: Using a balanced design provides ERP evidence against "syntax-first" approaches to sentence processing.

In this event-related potential (ERP) study we reevaluate syntax-first approaches to sentence processing by implementing a novel paradigm in French that includes correct sentences, pure syntactic category violations, lexical-semantic anomalies, and combined anomalies. Our balanced design systematically controlled for target word (noun vs. verb) and the context immediately preceding it. Group results from 36 native speakers of Quebec French revealed that, up to 300 ms, ERPs elicited by syntactic category violations were comparable with ERP responses to correct sentences, showing that there is no early activation reflecting syntactic category identification. Instead, in response to all anomalous conditions, we observed an N400 followed by a P600. Combined anomalies yielded additive effects of syntactic category and lexical-semantic anomalies on the N400, and a large P600 effect similar to the one observed in the pure syntactic condition. These results provide strong evidence against the hypothesis that (i) syntactic categories are processed first, and (ii) that syntactic category errors "block" lexical-semantic processing. Further, the N400 effect in response to pure syntactic category violations reflects a mismatch detection between a predicted word-stem and the actual target. This mechanism takes place simultaneously (and potentially in parallel) with lexical-semantic processing. In contrast, an interaction of syntax and semantics for the P600 reveals that the same neurocognitive resources are recruited for syntactic and semantic integration, both promoted by the implementation of an acceptability judgement task in our design. Additional analyses of individual data complemented these observations: during sentence processing, participants did not rely on one single cognitive mechanism reflected by either the N400 or the P600 effect but on both, suggesting that the biphasic N400-P600 ERP wave can indeed be considered to be an index of phrase-structure violations in most individuals, at least if they are realized on content words.