Working Memory Maintenance of Visual and Auditory Spatial Information Relies on Supramodal Neural Codes in the Dorsal Frontoparietal Cortex.

The frontoparietal attention network plays a pivotal role during working memory (WM) maintenance, especially under high-load conditions. Nevertheless, there is ongoing debate regarding whether this network relies on supramodal or modality-specific neural signatures. In this study, we used multi-voxel pattern analysis (MVPA) to evaluate the neural representation of visual versus auditory information during WM maintenance. During fMRI scanning, participants maintained small or large spatial configurations (low- or high-load trials) of either colour shades or sound pitches in WM for later retrieval. Participants were less accurate in retrieving high- vs. low-load trials, demonstrating an effective manipulation of WM load, irrespective of the sensory modality. The frontoparietal regions involved in maintaining high- vs. low-load spatial maps in either sensory modality were highlighted using a conjunction analysis. Widespread activity was found across the dorsal frontoparietal network, peaking on the frontal eye fields and the superior parietal lobule, bilaterally. Within these regions, MVPAs were performed to quantify the pattern of distinctness of visual vs. auditory neural codes during WM maintenance. These analyses failed to reveal distinguishable patterns in the dorsal frontoparietal regions, thus providing support for a common, supramodal neural code associated with the retention of either visual or auditory spatial configurations.

Consumer Neuroscience: Attentional Preferences for Wine Labeling Reflected in the Posterior Contralateral Negativity.

During the decision-making process, consumers notice, inspect, and visually scan different products. External characteristics of a product, such as design, packaging, label, and logo, have been shown to strongly influence how customers perceive, assess, and select a product. Marketers have put a lot of effort into determining the factors that trigger consumers' visual attention toward products, using traditional research methods, self-reports, or observations. The use of neuroscientific tools to study consumer behavior may improve our understanding of how external characteristics influence consumers' visual attention. Consumer neuroscience research shows that preferences for a product may already be reflected in brain activity before customers make a final decision. Using electroencephalography (EEG), we investigated whether the design of different wine labeling influences individual preferences, reflected in the neural activity related to visual attention. More specifically, we examined whether the posterior contralateral negativity (PCN) can be used to assess and predict consumers' preferences for a specific product based on its external characteristics. The PCN is commonly used to estimate attentional selection by focusing on stimulus-side dependent EEG lateralization above parieto-occipital areas. We computed the PCN to assess whether a certain wine label caught participants' visual attention and additionally by comparing the PCN with behavioral data (wine preferences and reaction times) to determine whether early effects of visual attention could predict participants' final preferences for a specific label. Our findings indicate that the PCN provides relevant information on visual attention mechanisms for external characteristics, as the view of the four labels modulated PCN amplitude. We hope this study can help researchers and practitioners in examining the effects of external product characteristics on consumer choice by estimating the changes in the EEG that are related to visual attention.

Frontal brain areas are more involved during motor imagery than during motor execution/preparation of a response sequence.

Results of several neuroimaging studies support the functional equivalence model, which states that motor imagery (MI) and motor execution (ME) involve the same processes, except for the final execution component. In contrast, the motor-cognitive model implies that MI additionally involves frontal executive control processes. However, according to some authors MI may actually be more comparable to motor preparation (MP). In the current electroencephalographic study, a version of the discrete sequence production paradigm was employed in which human participants initially had to prepare a sequence of five finger movements that subsequently had to be executed, imagined, or withheld. MI, ME, and MP were compared by computing event-related (de)-synchronization in the theta, alpha/mu, and beta bands. Results revealed a major increase in frontal theta power during MI as compared to ME and MP. At the end of the examined intervals, a posterior reduction in alpha power was present during ME and MP, but not during MI. Finally, above sensorimotor areas a decrease in beta power was observed that was most pronounced in the case of ME. The increase of frontal theta activity during MI may reflect increased effort, while the absence of a reduction in posterior alpha power suggests no major involvement of visuospatial attention and/or visual imagery. The present findings favor the motor-cognitive model, as it predicts extra involvement of frontal executive processes during MI.

The Simon effect in a discrete sequence production task: Key-specific stimuli cannot be ignored due to attentional capture.

Two experiments examined whether practicing discrete key pressing sequences eventually leads to a disregard of the key-specific stimuli, as suggested by sequence learning models, or whether these stimuli continue to be relied upon because the associated luminance increase attracts visuospatial attention. Participants practiced two sequences by reacting to two fixed series of seven letter stimuli, each displayed at a location that did or did not correspond with the required response location. Stimulus use was indicated by a Simon effect in that key presses were slowed when stimulus and key locations did not correspond. Experiment 1 demonstrated that letter stimuli continued to be used as the Simon effect occurred with each sequence element, and this remained quite stable across practice and did not differ for familiar and unfamiliar sequences. Experiment 2 showed that the Simon effect remained present even with meaningless stimuli that were often even harmful. These findings suggest that even in motor sequences that can be executed without element-specific stimuli attention attraction enforces stimulus use. The data further supported the assumptions that S-R translation and sequencing systems are racing to trigger individual responses, and that explicit sequence representations include spatial and verbal knowledge.

Does the influence of near-threshold primes depend on the type of task?

Most studies investigating the influence of primes on the processing of subsequent targets involve a main task in which responses are made to the targets, and a task that tests prime awareness. If the participant is not aware of the prime location/identity but an influence of the prime is observed in the main task, researchers conclude that this influence can be ascribed to unconscious processing of the prime. This implies the assumption that the prime's influence is independent of task instructions: a prime consciously perceived in the prime task is consciously perceived in the main task. In the metacontrast-masking study, we compared motor- and attention-related electroencephalographic (EEG) components in three tasks with the same stimuli but different instructions and showed that early posterior contralateral negativities (PCNs) and lateralized readiness potentials (LRPs) were smaller when primes were task-relevant than when targets were task-relevant. This suggests that early components may depend on task instruction and are not purely prime-related.

Dyslexic individuals orient but do not sustain visual attention: Electrophysiological support from the lower and upper alpha bands.

Individuals with developmental dyslexia have been characterized by problems with attentional orienting. In the current study, we specifically focused on possible changes in endogenous visual orienting that may be reflected in the electroencephalogram. A variant of the Posner cuing paradigm was employed with valid or invalid central cues that preceded target stimuli that were presented in the left or right visual field. The target stimuli consisted of vertical or horizontal stripes with low (two thick lines) or high (six thin lines) spatial frequencies. We examined lateralized alpha power in the cue-target interval as recent studies revealed that a contra vs. ipsilateral reduction in alpha power relates to the orienting of attention. An initial orienting effect in the lower alpha band was more pronounced for dyslexic individuals than for controls, suggesting that they oriented at an earlier moment in time. However, in contrast with controls, at the end of the cue-target interval no clear contralateral reduction in the upper alpha band was observed for dyslexic individuals. Dyslexic individuals additionally displayed slower responses, especially for invalidly cued high spatial frequency targets in the left visual field. The current data support the view that dyslexic individuals orient well to the cued location but have a problem with sustaining their attention.

Does Transcranial Direct Current Stimulation Affect the Learning of a Fine Sequential Hand Motor Skill with Motor Imagery?

Learning a fine sequential hand motor skill, like playing the piano or learning to type, improves not only due to physical practice, but also due to motor imagery. Previous studies revealed that transcranial direct current stimulation (tDCS) and motor imagery independently affect motor learning. In the present study, we investigated whether tDCS combined with motor imagery above the primary motor cortex influences sequence-specific learning. Four groups of participants were involved: an anodal, cathodal, sham stimulation, and a control group (without stimulation). A modified discrete sequence production (DSP) task was employed: the Go/NoGo DSP task. After a sequence of spatial cues, a response sequence had to be either executed, imagined, or withheld. This task allows to estimate both non-specific learning and sequence-specific learning effects by comparing the execution of unfamiliar sequences, familiar imagined, familiar withheld, and familiar executed sequences in a test phase. Results showed that the effects of anodal tDCS were already developing during the practice phase, while no effects of tDCS on sequence-specific learning were visible during the test phase. Results clearly showed that motor imagery itself influences sequence learning, but we also revealed that tDCS does not increase the influence of motor imagery on sequence learning.

Do musicians learn a fine sequential hand motor skill differently than non-musicians?

Do professional musicians learn a fine sequential hand motor skill more efficiently than non-musicians? Is this also the case when they perform motor imagery, which implies that they only mentally simulate these movements? Musicians and non-musicians performed a Go/NoGo discrete sequence production (DSP) task, which allows to separate sequence-specific from a-specific learning effects. In this task five stimuli, to be memorized during a preparation interval, signaled a response sequence. In a practice phase, different response sequences had to be either executed, imagined, or inhibited, which was indicated by different response cues. In a test phase, responses were required to familiar (previously executed, imagined, or inhibited) and unfamiliar sequences. In both phases, response times and response accuracy were measured while the electroencephalogram (EEG) was only registered during the practice phase to compare activity between motor imagery, motor execution, and motor inhibition for both groups. Results in the practice phase revealed that musicians learned the response sequences faster and more accurately than non-musicians although no difference in initiation time was found. EEG analyses revealed similar lateralized activity during learning a motor skill for both groups. Our results from the test phase showed better sequence-a-specific learning effects (i.e., faster response times and increased accuracy) for musicians than for non-musicians. Moreover, we revealed that non-musicians benefit more from physical execution while learning a required motor sequence, whereas sequence-specific learning effects due to learning with motor imagery were very similar for musicians and non-musicians.

Does mindfulness training modulate the influence of spatial attention on the processing of intracutaneous electrical stimuli?

Mindfulness based stress reduction (MBSR) training has been proposed to improve attentional skills by modulating thalamo-cortical loops that affect the sensitivity of relevant cortical areas like the somatosensory cortex. This modulation may be reflected in the electroencephalographic (EEG) alpha rhythm, and could affect the processing of subsequently applied intracutaneous electrical stimuli. Participants took part in an MBSR training and participated in two EEG sessions. EEG was measured in variants of an endogenous orienting paradigm in which attention had to be directed to the left or right forearm. After the orienting interval, the electrical stimulus was applied, equally likely on the attended or the unattended forearm. One group of participants took part in the EEG session before and after the training, while the other group took part after the training, and another time, eight weeks later. The influence of the MBSR training and spatial attention were examined with behavioral measures, lateralized alpha power within the orienting interval, and with event-related potentials (ERPs) evoked by the electrical stimuli. Self-reported mindfulness was clearly affected by the training, but no influence was found on other behavioral measures. Alpha power was clearly lateralized due to spatial attention and several ERP components (N130, N180, P340) were modulated by spatial attention but no support was found for an influence of the MBSR training. Finally, analyses revealed that individual differences in training time modulated some of the observed effects, but no support was found for an influence on attentional orienting.

The engagement of cortical areas preceding exogenous vergence eye movements.

Source analyses on event related potentials (ERPs) derived from the electroencephalogram (EEG) were performed to examine the respective roles of cortical areas preceding exogenously triggered saccades, combined convergences, and combined divergences. All eye movements were triggered by the offset of a central fixation light emitting diode (LED) and the onset of a lateral LED at various depths in an otherwise fully darkened room. Our analyses revealed that three source pairs, two located in the frontal lobe-the frontal eye fields (FEF) and an anterior frontal area-, and one located within the occipital cortex, can account for 99.2% of the observed ERPs. Overall, the comparison between source activities revealed the largest activity in the occipital cortex, while no difference in activity between FEF and the anterior frontal area was obtained. For all sources, increased activity was observed for combined vergences, especially combined convergences, relative to saccades. Behavioral results revealed that onset latencies were longest for combined convergences, intermediate for combined divergences, and the shortest for saccades. Together, these findings fit within a perspective in which both occipital and frontal areas play an important role in retinal disparity detection. In the case of saccades and combined divergences stimulus-locked activity was larger than response-locked activity, while no difference between stimulus- and response-locked activity was observed for combined convergences. These findings seem to imply that the electrophysiological activity preceding exogenous eye movements consists of a sensory-related part that is under cortical control, while subcortical structures may be held responsible for final execution.

On understanding creative language: The late positive complex and novel metaphor comprehension.

Novel metaphoric sentences have repeatedly evoked larger N400 amplitudes than literal sentences, while investigations of the late positive complex (LPC) have brought inconsistent results, with reports of both increased and reduced amplitudes. In two experiments, we examined novel metaphor comprehension in Polish, using the same set of literal, novel metaphoric, and anomalous sentences. The first aim of the study was to test whether novel metaphors would evoke larger or smaller late positivity complex (LPC) amplitudes compared to literal and anomalous sentences. Some earlier studies have shown that whether increased LPC amplitudes are observed or not may be related to the task participants are asked to perform, with explicit acceptability judgments being more likely to evoke the LPC effect. The second aim of the study was, thus, to test whether the observed LPC pattern would be the same across two tasks, the semantic decision task (Experiment 1) and the reading task (Experiment 2). Our results replicated the N400 effect observed in earlier studies on metaphor in both experiments. Most importantly, a reduction in late positivity to novel metaphors relative to anomalous sentences was found in both experiments. Additionally, this difference was broadly distributed over parietal sites in Experiment 1, and clearly left-lateralized in Experiment 2, which might imply differences in the involvement of recollection and semantic processes. Overall, these results seem to indicate that both conventionality and task demands modulate the LPC pattern.

How effector-specific is the effect of sequence learning by motor execution and motor imagery?

The aim of the present study was twofold. First, we wanted to examine how effector specific the effect of sequence learning by motor execution is, and second, we wanted to compare this effect with learning by motor imagery. We employed a Go/NoGo discrete sequence production task in which in each trial a spatial sequence of five stimuli was presented. After a Go signal the corresponding spatial response sequence had to be executed, while after a NoGo signal, the response sequence had to be mentally imagined. For the training phase, participants were divided into two groups. In the index finger group, participants had to respond (physically or mentally) with the left or right index finger, while in the hand group they had to respond with four fingers of the left or right hand. In a final test phase both execution modes were compared and all trials had to be executed. Response times and the percentage of correct responses were determined to establish learning effects. Results showed that sequence learning effects as assessed in the test phase were independent of the effector used during the training phase. Results revealed the presence of aspecific learning effects in the case of learning a required motor task with an index finger, but sequence-specific learning effects, both due to motor execution and to motor imagery, were not effector specific.

Endogenous spatial attention directed to intracutaneous electrical stimuli on the forearms involves an external reference frame.

In the present study, we examined whether the direction of attention while anticipating intracutaneous electrical stimuli on the left or right forearm occurs within an internal somatotopic or an external body-based reference frame. Participants placed their hands on a table in front of them in a normal position or in a crossed-hands position. A symbolic cue with a validity of 80% instructed participants to attend to either the left or the right side, which varied from trial to trial. Crossing the hands induces a conflict of internal and external reference frames which allows to determine the dominating reference frame(s). Analyses of the electroencephalogram (EEG) during the orienting phase revealed that crossing the arms did not induce a reversal of neural activity over central sites as a late direction attention-related positivity and increased ipsilateral alpha power over occipital and central sites was observed in both conditions. Hand position influenced the processing of the electrical stimuli as no effect of cue validity was observed on the P3a component in the crossed-hands position. Our results indicate that endogenous spatial attention to intracutaneous electrical stimuli primarily occurs within an external reference frame.

Comparing the effects of sustained and transient spatial attention on the orienting towards and the processing of electrical nociceptive stimuli.

We examined whether sustained vs. transient spatial attention differentially affect the processing of electrical nociceptive stimuli. Cued nociceptive stimuli of a relevant intensity (low or high) on the left or right forearm required a foot pedal press. The cued side varied trial wise in the transient attention condition, while it remained constant during a series of trials in the sustained attention condition. The orienting phase preceding the nociceptive stimuli was examined by focusing on lateralized EEG activity. ERPs were computed to examine the influence of spatial attention on the processing of the nociceptive stimuli. Results for the orienting phase showed increased ipsilateral alpha and beta power above somatosensory areas in both the transient and the sustained attention conditions, which may reflect inhibition of ipsilateral and/or disinhibition of contralateral somatosensory areas. Cued nociceptive stimuli evoked a larger N130 than uncued stimuli, both in the transient and the sustained attention conditions. Support for increased efficiency of spatial attention in the sustained attention condition was obtained for the N180 and the P540 component. We concluded that spatial attention is more efficient in the case of sustained than in the case of transient spatial attention.

Does sensitivity in binary choice tasks depend on response modality?

In most models of vision, a stimulus is processed in a series of dedicated visual areas, leading to categorization of this stimulus, and possible decision, which subsequently may be mapped onto a motor-response. In these models, stimulus processing is thought to be independent of the response modality. However, in theories of event coding, common coding, and sensorimotor contingency, stimuli may be very specifically mapped onto certain motor-responses. Here, we compared performance in a shape localization task and used three different response modalities: manual, saccadic, and verbal. Meta-contrast masking was employed at various inter-stimulus intervals (ISI) to manipulate target visibility. Although we found major differences in reaction times for the three response modalities, accuracy remained at the same level for each response modality (and all ISIs). Our results support the view that stimulus-response (S-R) associations exist only for specific instances, such as reflexes or skills, but not for arbitrary S-R pairings.

Two Sides of the Same Coin: ERP and Wavelet Analyses of Visual Potentials Evoked and Induced by Task-Relevant Faces.

New analysis techniques of the electroencephalogram (EEG) such as wavelet analysis open the possibility to address questions that may largely improve our understanding of the EEG and clarify its relation with related potentials (ER Ps). Three issues were addressed. 1) To what extent can early ERERP components be described as transient evoked oscillations in specific frequency bands? 2) Total EEG power (TP) after a stimulus consists of pre-stimulus baseline power (BP), evoked power (EP), and induced power (IP), but what are their respective contributions? 3) The Phase Reset model proposes that BP predicts EP, while the evoked model holds that BP is unrelated to EP; which model is the most valid one? EEG results on NoGo trials for 123 individuals that took part in an experiment with emotional facial expressions were examined by computing ERPs and by performing wavelet analyses on the raw EEG and on ER Ps. After performing several multiple regression analyses, we obtained the following answers. First, the P1, N1, and P2 components can by and large be described as transient oscillations in the α and θ bands. Secondly, it appears possible to estimate the separate contributions of EP, BP, and IP to TP, and importantly, the contribution of IP is mostly larger than that of EP. Finally, no strong support was obtained for either the Phase Reset or the Evoked model. Recent models are discussed that may better explain the relation between raw EEG and ERPs.

Editorial Special Issue: Neuronus.

This special issue of the 12th volume of Advances in Cognitive Psychology is devoted to the Neuronus conference that took place in Kraków in 2015. In this editorial letter, we will focus on a selection of the materials and some follow-up research that was presented during this conference. We will also briefly introduce the conference contributions that successfully passed an external reviewing process.

Children with unilateral cerebral palsy show diminished implicit motor imagery with the affected hand.

AIM: Motor imagery refers to the mental simulation of a motor action without producing an overt movement. Implicit motor imagery can be regarded as a first-person kinesthetic perceptual judgement, and addresses the capacity to engage into the manipulation of one's body schema. In this study, we examined whether children with unilateral cerebral palsy (CP) are able to engage in implicit motor imagery. METHOD: A modified version of the hand laterality judgment task was employed. Erroneous responses, reaction times, and event-related potentials from the electroencephalograph were analysed. RESULTS: In 13 children with typical development (mean age 10y 7mo, SD 1y 2mo; seven male, six female), we observed the classic rotation direction effect. Specifically, when comparing outward rotated with inward rotated hand pictures, decreased accuracy and increased response times were observed. Event-related potentials analyses of the electroencephalogram revealed a more marked N1 and an enhanced rotation-related negativity. INTERPRETATION: These findings suggest that an implicit motor imagery strategy was used to solve the task. However, in 10 children with unilateral CP (mean age 10y 7mo, SD 2y 5mo; five male, five female), these effects were observed only when the less-affected hand was involved. This observation suggests that children with CP could benefit from visual training strategies.

Internal and external spatial attention examined with lateralized EEG power spectra.

Several authors argued that retrieval of an item from visual short term memory (internal spatial attention) and focusing attention on an externally presented item (external spatial attention) are similar. Part of the neuroimaging support for this view may be due to the employed experimental procedures. Furthermore, as internal spatial attention may have a more induced than evoked nature some effects may not have been visible in event related analyses of the electroencephalogram (EEG), which limits the possibility to demonstrate differences. In the current study, a colored frame cued which stimulus, one out of four presented in separate quadrants, required a response, which depended on the form of the cued stimulus (circle or square). Importantly, the frame occurred either before (precue), simultaneously with (simultaneous cue), or after the stimuli (postcue). The precue and simultaneous cue condition both concern external attention, while the postcue condition implies the involvement of internal spatial attention. Event-related lateralizations (ERLs), reflecting evoked effects, and lateralized power spectra (LPS), reflecting both evoked and induced effects, were determined. ERLs revealed a posterior contralateral negativity (PCN) only in the precue condition. LPS analyses on the raw EEG showed early increased contralateral theta power at posterior sites and later increased ipsilateral alpha power at occipito-temporal sites in all cue conditions. Responses were faster when the internally or externally attended location corresponded with the required response side than when not. These findings provide further support for the view that internal and external spatial attention share their underlying mechanism.

Effects of hand orientation on motor imagery--event related potentials suggest kinesthetic motor imagery to solve the hand laterality judgment task.

Motor imagery (MI) refers to the process of imagining the execution of a specific motor action without actually producing an overt movement. Two forms of MI have been distinguished: visual MI and kinesthetic MI. To distinguish between these forms of MI we employed an event related potential (ERP) study to measure interference effects induced by hand orientation manipulations in a hand laterality judgement task. We hypothesized that this manipulation should only affect kinesthetic MI but not visual MI. The ERPs elicited by rotated hand stimuli contained the classic rotation related negativity (RRN) with respect to palm view stimuli. We observed that laterally rotated stimuli led to a more marked RRN than medially rotated stimuli. This RRN effect was observed when participants had their hands positioned in either a straight (control) or an inward rotated posture, but not when their hands were positioned in an outward rotated posture. Posture effects on the ERP-RRN have not previously been studied. Apparently, a congruent hand posture (hands positioned in an outward rotated fashion) facilitates the judgement of the otherwise more demanding laterally rotated hand stimuli. These ERP findings support a kinesthetic interpretation of MI involved in solving the hand laterality judgement task. The RRN may be used as a non-invasive marker for kinesthetic MI and seems useful in revealing the covert behavior of MI in e.g. rehabilitation programs.