Response stopping under conflict: The integrative role of representational dynamics associated with the insular cortex.

Coping with distracting inputs during goal-directed behavior is a common challenge, especially when stopping ongoing responses. The neural basis for this remains debated. Our study explores this using a conflict-modulation Stop Signal task, integrating group independent component analysis (group-ICA), multivariate pattern analysis (MVPA), and EEG source localization analysis. Consistent with previous findings, we show that stopping performance is better in congruent (nonconflicting) trials than in incongruent (conflicting) trials. Conflict effects in incongruent trials compromise stopping more due to the need for the reconfiguration of stimulus-response (S-R) mappings. These cognitive dynamics are reflected by four independent neural activity patterns (ICA), each coding representational content (MVPA). It is shown that each component was equally important in predicting behavioral outcomes. The data support an emerging idea that perception-action integration in action-stopping involves multiple independent neural activity patterns. One pattern relates to the precuneus (BA 7) and is involved in attention and early S-R processes. Of note, three other independent neural activity patterns were associated with the insular cortex (BA13) in distinct time windows. These patterns reflect a role in early attentional selection but also show the reiterated processing of representational content relevant for stopping in different S-R mapping contexts. Moreover, the insular cortex's role in automatic versus complex response selection in relation to stopping processes is shown. Overall, the insular cortex is depicted as a brain hub, crucial for response selection and cancellation across both straightforward (automatic) and complex (conditional) S-R mappings, providing a neural basis for general cognitive accounts on action control.

The role of visual association cortices during response selection processes in interference-modulated response stopping.

Response inhibition and the ability to navigate distracting information are both integral parts of cognitive control and are imperative to adaptive behavior in everyday life. Thus far, research has only inconclusively been able to draw inferences regarding the association between response stopping and the effects of interfering information. Using a novel combination of the Simon task and a stop signal task, the current study set out to investigate the behavioral as well as the neurophysiological underpinnings of the relationship between response stopping and interference processing. We tested n = 27 healthy individuals and combined temporal EEG signal decomposition with source localization methods to delineate the precise neurophysiological dynamics and functional neuroanatomical structures associated with conflict effects on response stopping. The results showed that stopping performance was compromised by conflicts. Importantly, these behavioral effects were reflected by specific aspects of information coded in the neurophysiological signal, indicating that conflict effects during response stopping are not mediated via purely perceptual processes. Rather, it is the processing of specific, stop-relevant stimulus features in the sensory regions during response selection, which underlies the emergence of conflict effects in response stopping. The findings connect research regarding response stopping with overarching theoretical frameworks of perception-action integration.

Alcohol-induced deficits in reactive control of response selection and inhibition are counteracted by a seemingly paradox increase in proactive control.

High-dose alcohol intoxication reduces cognitive control, including inhibition. Although inhibition deficits may contribute to the behavioral deficits commonly observed in alcohol use disorder (AUD), many questions about potentially modulating factors have remained unanswered. We examined the effects of experimentally induced high-dose alcohol intoxication (~ 1.1 ‰) on the interplay between controlled vs. automatic response selection and inhibition in healthy young men. A holistic EEG-based theta activity analysis that considered both reactive control during task performance and preceding proactive control processes was run. It revealed a previously unknown seesaw relationship, with decreased reactive control, but paradoxically increased proactive control. Most importantly, alcohol-induced increases in proactive occipital theta band power were associated with reductions in negative alcohol effects on reactive control processes associated with decreased activity in the SMA and medial frontal cortex. Our findings demonstrate that research should not solely focus on immediate effects during task performance. Aside from differential neurobiochemical and neuroanatomical effects of alcohol, it is also conceivable that proactive control may have been recruited in a (secondary) response to compensate for alcohol-induced impairments in reactive control. Against this background, it could be promising to investigate changes in such compensatory mechanisms in pronounced alcohol-associated inhibition deficits, like in AUD patients.

Evidence for independent representational contents in inhibitory control subprocesses associated with frontoparietal cortices.

Inhibitory control processes have intensively been studied in cognitive science for the past decades. Even though the neural dynamics underlying these processes are increasingly better understood, a critical open question is how the representational dynamics of the inhibitory control processes are modulated when engaging in response inhibition in a relatively automatic or a controlled mode. Against the background of an overarching theory of perception-action integration, we combine temporal and spatial EEG signal decomposition methods with multivariate pattern analysis and source localization to obtain fine-grained insights into the neural dynamics of the representational content of response inhibition. For this purpose, we used a sample of N = 40 healthy adult participants. The behavioural data suggest that response inhibition was better in a more controlled than a more automated response execution mode. Regarding neural dynamics, effects of response inhibition modes relied on a concomitant coding of stimulus-related information and rules of how stimulus information is related to the appropriate motor programme. Crucially, these fractions of information, which are encoded at the same time in the neurophysiological signal, are based on two independent spatial neurophysiological activity patterns, also showing differences in the temporal stability of the representational content. Source localizations revealed that the precuneus and inferior parietal cortex regions are more relevant than prefrontal areas for the representation of stimulus-response selection codes. We provide a blueprint how a concatenation of EEG signal analysis methods, capturing distinct aspects of neural dynamics, can be connected to cognitive science theory on the importance of representations in action control.

On the Role of Stimulus-Response Context in Inhibitory Control in Alcohol Use Disorder.

The behavioral and neural dynamics of response inhibition deficits in alcohol use disorder (AUD) are still largely unclear, despite them possibly being key to the mechanistic understanding of the disorder. Our study investigated the effect of automatic vs. controlled processing during response inhibition in participants with mild-to-moderate AUD and matched healthy controls. For this, a Simon Nogo task was combined with EEG signal decomposition, multivariate pattern analysis (MVPA), and source localization methods. The final sample comprised n = 59 (32♂) AUD participants and n = 64 (28♂) control participants. Compared with the control group, AUD participants showed overall better response inhibition performance. Furthermore, the AUD group was less influenced by the modulatory effect of automatic vs. controlled processes during response inhibition (i.e., had a smaller Simon Nogo effect). The neurophysiological data revealed that the reduced Simon Nogo effect in the AUD group was associated with reduced activation differences between congruent and incongruent Nogo trials in the inferior and middle frontal gyrus. Notably, the drinking frequency (but not the number of AUD criteria we had used to distinguish groups) predicted the extent of the Simon Nogo effect. We suggest that the counterintuitive advantage of participants with mild-to-moderate AUD over those in the control group could be explained by the allostatic model of drinking effects.

The importance of resource allocation for the interplay between automatic and cognitive control in response inhibition - An EEG source localization study.

Goal-directed behavior often requires the inhibition of prepotent automatic responses. Response inhibition encompasses several top-down cognitive operations embedded in a neural network extending from fronto-cortical regions to subcortical nuclei. Yet, it has remained unclear whether the early allocation of cognitive resources also modulates response inhibition performance and neural structures involved in this process. To investigate this question, we used a Simon Nogo task, which was designed to manipulate the relationship between automaticity and cognitive control during response inhibition, and combined it with an electroencephalogram (EEG) and source localization approach. We showed that the early allocation of cognitive resources, as reflected by the P2 amplitude, might be a critical determinant in the interplay between automaticity and cognitive control in response inhibition. Specifically, the obtained results demonstrated that individual variations in cognitive resource allocation modulated the need for conflict monitoring and engagement in cognitive control processes, as reflected by N2 and P3b amplitudes, respectively. Importantly, larger P2 amplitudes were associated with higher activation in cortical regions encompassing the temporo-parietal junction (TPJ). This stresses the importance of this cortical region for the encoding of relevant stimulus information to resolve conflicting contexts in response inhibition. The increased cognitive control in more automatic contexts was also reflected by higher activation of the superior and medial frontal cortices. These findings provide a new perspective on response inhibition, suggesting that early resource allocation is a central modulator of the interaction between automaticity and cognitive control.

Auricular transcutaneous vagus nerve stimulation for alcohol use disorder: A chance to improve treatment?

Alcohol use disorder (AUD) is a relapsing-remitting condition characterized by excessive and/or continued alcohol consumption despite harmful consequences. New adjuvant tools, such as noninvasive brain stimulation techniques, might be helpful additions to conventional treatment approaches or even provide an alternative option for patients who fail to respond adequately to other treatment options. Here, we discuss the potential use of auricular transcutaneous vagus nerve stimulation (atVNS) as an ADD-ON intervention in AUD. Compared with other techniques, atVNS has the advantage of directly stimulating nuclei that synthesize GABA and catecholamines, both of which are functionally altered by alcohol intake in AUD patients. Pharmacological options targeting those neurotransmitters are widely available, but have relatively limited beneficial effects on cognition, even though restoring normal cognitive functioning, especially cognitive control, is key to maintaining abstinence. Against this background, atVNS could be a particularly useful add-on because there is substantial meta-analytic evidence based on studies in healthy individuals that atVNS can enhance cognitive control processes that are crucial to regaining control over drug intake. We discuss essential future research on using atVNS as an ADD-ON intervention in AUD to enhance clinical and cognitive outcomes by providing a translational application. Given that this novel technique can be worn like an earpiece and can be employed without medical supervision/outside the clinical settings, atVNS could be well integratable into the daily life of the patients, where the task of regaining control over drug intake is most challenging.

How low working memory demands and reduced anticipatory attentional gating contribute to impaired inhibition during acute alcohol intoxication.

High-dose alcohol intoxication is commonly associated with impaired inhibition, but the boundary conditions, as well as associated neurocognitive/neuroanatomical changes have remained rather unclear. This study was motivated by the counterintuitive finding that high-dose alcohol intoxication compromises response inhibition performance when working memory demands were low, but not when they were high. To investigate whether this is more likely to be caused by deficits in cognitive control processes or in attentional processes, we examined event-related (de)synchronization processes in theta and alpha-band activity and performed beamforming analyses on the EEG data of previously published behavioral findings. This yielded two possible explanations: There may be a selective decrease of working memory engagement in case of relatively low demand, which boosts response automatization, ultimately putting more strain on the remaining inhibitory resources. Alternatively, there may be a decrease in proactive preparatory and anticipatory attentional gating processes in case of relatively low demand, hindering attentional sampling of upcoming stimuli. Crucially, both of these interrelated mechanisms reflect differential alcohol effects after the actual motor inhibition process and therefore tend to be processes that serve to anticipate future response inhibition affordances. This provides new insights into how high-dose alcohol intoxication can impair inhibitory control.

Pretrial Theta Band Activity Affects Context-dependent Modulation of Response Inhibition.

The ability to inhibit a prepotent response is a crucial prerequisite of goal-directed behavior. So far, research on response inhibition has mainly examined these processes when there is little to no cognitive control during the decision to respond. We manipulated the "context" in which response inhibition has to be exerted (i.e., a controlled or an automated context) by combining a Simon task with a go/no-go task and focused on theta band activity. To investigate the role of "context" in response inhibition, we also examined how far theta band activity in the pretrial period modulates context-dependent variations of theta band activity during response inhibition. This was done in an EEG study applying beamforming methods. Here, we examined n = 43 individuals. We show that an automated context, as opposed to a controlled context, compromises response inhibition performance and increases the need for cognitive control. This was also related to context-dependent modulations of theta band activity in superior frontal and middle frontal regions. Of note, results showed that theta band activity in the pretrial period, associated with the right inferior frontal cortex, was substantially correlated with context-dependent modulations of theta band activity during response inhibition. The direction of the obtained correlation provides insights into the functional relevance of a pretrial theta band activity. The data suggest that pretrial theta band activity reflects some form of attentional sampling to inform possible upcoming processes signaling the need for cognitive control.

A role of the norepinephrine system or effort in the interplay of different facets of inhibitory control.

Inhibitory control has multiple facets, and one possible distinction can be made between 'inhibition of interferences' and the 'inhibition of actions'. Both facets of inhibitory control show an interdependency. Even though some neurophysiological processes underlying this interdependency have been examined, the role of neuro-modulatory processes in their interplay are not understood. In the current study, we examine the role of the norepinephrine (NE) system in these processes. We did so by combining a Go/Nogo and Simon task. We recorded the EEG and pupil diameter data as an indirect index of NE system activity during the task. EEG theta band activity data and pupil diameter data were then integrated after conducting a temporal signal decomposition of the EEG data. We show that particularly theta band activity coding stimulus-response translation processes associated with middle frontal cortices, but not stimulus-driven processes are modulated by the interplay between the 'inhibition of interferences' and the 'inhibition of actions'. Modulations in stimulus-response translation processes were systematically correlated with pupil-diameter responses. The pattern of correlations suggests that phasic NE system activity particularly modulates stimulus-response mapping processes during conflict monitoring in incongruent Nogo trials, which may explain behavioral performance effects. Phasic NE system activity reflects essential modulators of the interplay between the 'inhibition of interferences' and the 'inhibition of actions'.

Neurobiological mechanisms of control in alcohol use disorder - Moving towards mechanism-based non-invasive brain stimulation treatments.

Alcohol use disorder (AUD) is characterized by excessive habitual drinking and loss of control over alcohol intake despite negative consequences. Both of these aspects foster uncontrolled drinking and high relapse rates in AUD patients. Yet, common interventions mostly focus on the phenomenological level, and prioritize the reduction of craving and withdrawal symptoms. Our review provides a mechanistic understanding of AUD and suggests alternative therapeutic approaches targeting the mechanisms underlying dysfunctional alcohol-related behaviours. Specifically, we explain how repeated drinking fosters the development of rigid drinking habits and is associated with diminished cognitive control. These behavioural and cognitive effects are then functionally related to the neurobiochemical effects of alcohol abuse. We further explain how alterations in fronto-striatal network activity may constitute the neurobiological correlates of these alcohol-related dysfunctions. Finally, we discuss limitations in current pharmacological AUD therapies and suggest non-invasive brain stimulation (like TMS and tDCS interventions) as a potential addition/alternative for modulating the activation of both cortical and subcortical areas to help re-establish the functional balance between controlled and automatic behaviour.

Visual Short-Term Memory for Coherent and Sequential Motion: A rTMS Investigation.

We investigated the role of the human medio-temporal complex (hMT+) in the memory encoding and storage of a sequence of four coherently moving random dot kinematograms (RDKs), by applying repetitive transcranial magnetic stimulation (rTMS) during an early or late phase of the retention interval. Moreover, in a second experiment, we also tested whether disrupting the functional integrity of hMT+ during the early phase impaired the precision of the encoded motion directions. Overall, results showed that both recognition accuracy and precision were worse in middle serial positions, suggesting the occurrence of primacy and recency effects. We found that rTMS delivered during the early (but not the late) phase of the retention interval was able to impair not only recognition of RDKs, but also the precision of the retained motion direction. However, such impairment occurred only for RDKs presented in middle positions along the presented sequence, where performance was already closer to chance level. Altogether these findings suggest an involvement of hMT+ in the memory encoding of visual motion direction. Given that both position sequence and rTMS modulated not only recognition but also the precision of the stored information, these findings are in support of a model of visual short-term memory with a variable resolution of each stored item, consistent with the assigned amount of memory resources, and that such item-specific memory resolution is supported by the functional integrity of area hMT+.

Spatial and Temporal Selectivity of Translational Glass Patterns Assessed With the Tilt After-Effect.

Glass patterns (GPs) have been widely employed to investigate the mechanisms underlying processing of global form from locally oriented cues. The current study aimed to psychophysically investigate the level at which global orientation is extracted from translational GPs using the tilt after-effect (TAE) and manipulating the spatiotemporal properties of the adapting pattern. We adapted participants to translational GPs and tested with sinewave gratings. In Experiment 1, we investigated whether orientation-selective units are sensitive to the temporal frequency of the adapting GP. We used static and dynamic translational GPs, with dynamic GPs refreshed at different temporal frequencies. In Experiment 2, we investigated the spatial frequency selectivity of orientation-selective units by manipulating the spatial frequency content of the adapting GPs. The results showed that the TAE peaked at a temporal frequency of ∼30 Hz, suggesting that orientation-selective units responding to translational GPs are sensitive to high temporal frequencies. In addition, TAE from translational GPs peaked at lower spatial frequencies than the dipoles' spatial constant. These effects are consistent with form-motion integration at low and intermediate levels of visual processing.

Electrophysiological aftereffects of high-frequency transcranial random noise stimulation (hf-tRNS): an EEG investigation.

There is evidence that high-frequency transcranial random noise stimulation (hf-tRNS) is effective in improving behavioural performance in several visual tasks. However, so far there has been limited research into the spatial and temporal characteristics of hf-tRNS-induced facilitatory effects. In the present study, electroencephalogram (EEG) was used to investigate the spatial and temporal dynamics of cortical activity modulated by offline hf-tRNS on performance on a motion direction discrimination task. We used EEG to measure the amplitude of motion-related VEPs over the parieto-occipital cortex, as well as oscillatory power spectral density (PSD) at rest. A time-frequency decomposition analysis was also performed to investigate the shift in event-related spectral perturbation (ERSP) in response to the motion stimuli between the pre- and post-stimulation period. The results showed that the accuracy of the motion direction discrimination task was not modulated by offline hf-tRNS. Although the motion task was able to elicit motion-dependent VEP components (P1, N2, and P2), none of them showed any significant change between pre- and post-stimulation. We also found a time-dependent increase of the PSD in alpha and beta bands regardless of the stimulation protocol. Finally, time-frequency analysis showed a modulation of ERSP power in the hf-tRNS condition for gamma activity when compared to pre-stimulation periods and Sham stimulation. Overall, these results show that offline hf-tRNS may induce moderate aftereffects in brain oscillatory activity.

Alcohol intoxication, but not hangover, differentially impairs learning and automatization of complex motor response sequences.

Behavioral automatization usually makes us more efficient and less error-prone, but may also foster dysfunctional behavior like alcohol abuse. Yet, it has remained unclear whether alcohol itself causes the shift from controlled to habitual behavior commonly observed in alcohol use disorder (AUD). We thus investigated how the acute and post-acute effects of binge drinking affect the automatization of motor response sequences and the execution of automated vs. controlled motor response sequences. N = 70 healthy young men performed a newly developed automatization paradigm once sober and once after binge drinking (half of them intoxicated and half of them hungover). While we found no significant effects of alcohol hangover, acute intoxication (~ 1.2 ‰) had two dissociable effects: Firstly, it impaired the automatization of complex motor response sequence execution. Secondly, it eliminated learning effects in response selection and pre-motor planning processes. The results suggest that alcohol hangover did not affect controlled or automated processes, and disprove the assumption that alcohol intoxication generally spares or facilitates motor response sequence automatization. As these effects could be specific to the investigated explicit learning context, acute intoxication might potentially still improve the execution of pre-existing automatisms and/or the implicit acquisition of motor response sequence automatisms.

Visual short-term memory for coherent motion in video game players: evidence from a memory-masking paradigm.

In this study, we investigated visual short-term memory for coherent motion in action video game players (AVGPs), non-action video game players (NAVGPs), and non-gamers (control group: CONs). Participants performed a visual memory-masking paradigm previously used with macaque monkeys and humans. In particular, we tested whether video game players form a more robust visual short-term memory trace for coherent moving stimuli during the encoding phase, and whether such memory traces are less affected by an intervening masking stimulus presented 0.2 s after the offset of the to-be-remembered sample. The results showed that task performance of all groups was affected by the masking stimulus, but video game players were affected to a lesser extent than controls. Modelling of performance values and reaction times revealed that video game players have a lower guessing rate than CONs, and higher drift rates than CONs, indicative of more efficient perceptual decisions. These results suggest that video game players exhibit a more robust VSTM trace for moving objects and this trace is less prone to external interference.

Limited Attention Diminishes Spatial Suppression From Large Field Glass Patterns.

Glass patterns (GPs) consist of randomly distributed dot pairs (dipoles) whose orientations are determined by specific geometric transforms. We investigated the role of visuospatial attention in the processing of global form from GPs by measuring the effect of distraction on adaptation to GPs. In the nondistracted condition, observers were adapted to coherent GPs. After the adaptation period, they were presented with a test GP divided in two halves along the vertical and were required to judge which side of the test GP was more coherent. In the attention-distracted condition, a high-load rapid serial visual presentation task was performed during the adapting period. The magnitude of the form after-effect was measured using a technique that measures the coherence level at which the test GP appears random. The rationale was that if attention has a modulatory effect on the spatial summation of dipoles, in the attention-distracted condition, we should expect a weaker form after-effect. However, the results showed stronger form after-effect in the attention-distracted condition than in the nondistracted condition, suggesting that distraction during adaptation increases the strength of form adaptation. Additional experiments suggested that distraction may reduce the spatial suppression from large-scale textures, strengthening the spatial summation of local-oriented signals.

Modulatory mechanisms underlying high-frequency transcranial random noise stimulation (hf-tRNS): A combined stochastic resonance and equivalent noise approach.

BACKGROUND: High-frequency transcranial random noise stimulation (hf-tRNS) is a neuromodulatory technique consisting of the application of alternating current at random intensities and frequencies. hf-tRNS induces random neural activity in the system that may boost the sensitivity of neurons to weak inputs. Stochastic resonance is a nonlinear phenomenon whereby the addition of an optimal amount of noise results in performance enhancement, whereas further noise increments impair signal detection or discrimination. OBJECTIVE: The aim of the study was to assess whether modulatory effects of hf-tRNS rely on the stochastic resonance phenomenon, and what is the specific neural mechanism producing stochastic resonance. METHOD: Observers performed a two-interval forced choice motion direction discrimination task in which they had to report whether two moving patches presented in two temporal intervals had the same or different motion directions. hf-tRNS was administered at five intensity levels (0.5, 0.75, 1.0, 1.5, and 2.25 mA). RESULTS: The results showed a significant improvement in performance when hf-tRNS was applied at 1.5 mA, representing the optimal level of external noise. However, stimulation intensity at 2.25 mA significantly impaired direction discrimination performance. An equivalent noise (EN) analysis, used to assess how hf-tRNS modulates the mechanisms underlying global motion processing, showed an increment in motion signal integration with the optimal current intensity, but reduced motion signal integration at 2.25 mA. CONCLUSION: These results indicate that hf-tRNS-induced noise modulates neural signal-to-noise ratio in a way that is compatible with the stochastic resonance phenomenon.

The effects of high-frequency transcranial random noise stimulation (hf-tRNS) on global motion processing: An equivalent noise approach.

BACKGROUND: High frequency transcranial random noise stimulation (hf-tRNS) facilitates performance in several perceptual and cognitive tasks, however, little is known about the underlying modulatory mechanisms. OBJECTIVE: In this study we compared the effects of hf-tRNS to those of anodal and cathodal tDCS in a global motion direction discrimination task. An equivalent noise (EN) paradigm was used to assess how hf-tRNS modulates the mechanisms underlying local and global motion processing. METHOD: Motion coherence threshold and slope of the psychometric function were estimated using an 8AFC task in which observers had to discriminate the motion direction of a random dot kinematogram presented either in the left or right visual hemi-field. During the task hf-tRNS, anodal and cathodal tDCS were delivered over the left hMT+. In a subsequent experiment we implemented an EN paradigm in order to investigate the effects of hf-tRNS on the mechanisms involved in visual motion integration (i.e., internal noise and sampling). RESULTS: hf-tRNS reduced the motion coherence threshold but did not affect the slope of the psychometric function, suggesting no modulation of stimulus discriminability. Anodal and cathodal tDCS did not produce any modulatory effects. EN analysis in the last experiment found that hf-tRNS modulates sampling but not internal noise, suggesting that hf-tRNS modulates the integration of local motion cues. CONCLUSION: hf-tRNS interacts with the output neurons tuned to directions near to the directional signal, incrementing the signal-to-noise ratio and the pooling of local motion cues and thus increasing the sensitivity for global moving stimuli.

The interaction between orientation and motion signals in moving oriented Glass patterns.

Previous psychophysical evidence suggests that motion and orientation processing systems interact asymmetrically in the human visual system, with orientation information having a stronger influence on the perceived motion direction than vice versa. To investigate the mechanisms underlying this motion-form interaction we used moving and oriented Glass patterns (GPs), which consist of randomly distributed dot pairs (dipoles) that induce the percept of an oriented texture. In Experiment 1 we varied the angle between dipole orientation and motion direction (conflict angle). In separate sessions participants either judged the orientation or motion direction of the GP. In addition, the spatiotemporal characteristics of dipole motion were manipulated as a way to limit (Experiment 1) or favor (Experiment 2) the availability of orientation signals from motion (motion streaks). The results of Experiment 1 showed that apparent GP motion direction is attracted toward dipole orientation, and apparent GP orientation is repulsed from GP motion. The results of Experiment 2 showed stronger repulsion effects when judging the GP orientation, but stronger motion streaks from the GP motion can dominate over the signals provided by conflicting dipole orientation. These results are consistent with the proposal that two separate mechanisms contribute to our perception of stimuli which contain conflicting orientation and motion information: (i) perceived GP motion is mediated by spatial motion-direction sensors, in which signals from motion sensors are combined with excitatory input from orientation-tuned sensors tuned to orientations parallel to the axis of GP motion, (ii) perceived GP orientation is mediated by orientation-tuned sensors which mutually inhibit each other. The two mechanisms are revealed by the different effects of conflict angle and dipole lifetime on perceived orientation and motion direction.