Biology subjects

Fahrenfort, J. J.

Publications and source records attributed to Fahrenfort, J. J..

At least 19 recordsLinked to original sources

bioRxiv2026

Goal-directed action selection relies on conjunctive representations that bind goals, actions and expected sensory outcomes

Goal-directed actions are performed to obtain specific sensory outcomes. However, theories of action control diverge on whether anticipated outcomes are only consequences of motor command specification or serve as instrumental determinants of action selection. Here, we directly tested whether predicted sensory outcomes are integrated into the neural processes underlying the selection of goal-directed actions. Participants learned novel action-outcome contingencies and selected actions to generate specific sensory outcomes, while their brain activity was recorded using EEG. Combining time-resolved multivariate decoding with representational similarity analysis, we tracked both feature-specific and conjunctive neural representations over time. We found that the strength of high-dimensional conjunctive representations including anticipated sensory outcomes predicted trial-to-trial reaction times, even after accounting for individual task features. These results provide neural evidence that predicted sensory outcomes are not downstream consequences, but core determinants of action selection, supporting the view that actions are selected to control expected sensory input.

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bioRxiv2026

Effects of expectation, attention, and NMDA receptor blockade on feedforward and feedback processing

Perception is increasingly viewed as an inferential process wherein sensory inputs are integrated with prior expectations. We employed time-resolved decoding on electroencephalography (EEG) data (n = 30 male participants) to investigate how expectations modulate sensory processing across varying levels of stimulus complexity, and tested the effect of attention and NMDA receptor blockade. We designed a visual stimulus containing features of different complexity whose processing relies on distinct neural mechanisms: local contrast, collinearity, and the Kanizsa illusion, involving primarily feedforward, lateral, and feedback processes, respectively. EEG decoding revealed that expectations modulated lateral and feedback processing (better decoding for unexpected stimuli) but not feedforward processing. These expectation effects were confined to attended (task-relevant) features and were not observed for task-irrelevant features. The NMDA receptor antagonist memantine selectively enhanced decoding of the Kanizsa illusion, implicating NMDA-mediated feedback mechanisms in perceptual inference, but it did not modulate the effects of expectation or attention. These findings highlight the differential impact of expectations across different stages of sensory processing and reveal a distinct role of NMDA receptor-mediated feedback mechanisms. Significance statementPerception integrates sensory inputs with prior expectations. Using EEG decoding, we examined how expectations shape sensory processing at different levels of complexity and tested the effects of attention and NMDA receptor blockade. Our visual stimuli were designed to capture EEG markers of feedforward, lateral, and feedback mechanisms. Expectations influenced lateral and feedback processing (better decoding for unexpected stimuli) but not feedforward processing, and these effects were selective to task-relevant stimulus features. Memantine, an NMDA receptor antagonist, selectively improved decoding of the Kanizsa illusion, implicating NMDA-mediated feedback in perceptual inference, but it did not shape expectation or attention effects.

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bioRxiv2024

Adaptive arousal regulation: Pharmacologically shifting the peak of the Yerkes-Dodson curve by catecholaminergic enhancement of arousal

Performance typically peaks at moderate arousal levels, consistent with the Yerkes-Dodson law, as confirmed by recent human and mouse pupillometry studies. Arousal states are influenced by neuromodulators like catecholamines (noradrenaline; NA and dopamine; DA) and acetylcholine (ACh). To explore their causal roles in this law, we pharmacologically enhanced arousal while measuring human decision-making and spontaneous arousal fluctuations via pupil size. The catecholaminergic agent atomoxetine (ATX) increased overall arousal and shifted the entire arousal-performance curve, suggesting a relative arousal mechanism where performance adapts to arousal fluctuations within arousal states. In contrast, the cholinergic agent donepezil (DNP) did not affect arousal or the curve. We modeled these findings in a neurobiologically plausible computational framework, showing how catecholaminergic modulation alters a disinhibitory neural circuit that encodes sensory evidence for decision-making. This work suggests that performance adapts flexibly to arousal fluctuations, ensuring optimal performance in each and every arousal state.

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bioRxiv2024

NMDA receptor antagonist memantine selectively affects recurrent processing during perceptual inference

Perceptual inference requires the integration of visual features through recurrent processing, the dynamic exchange of information between higher and lower level cortical regions. While animal research has demonstrated a crucial role of NMDA receptors in recurrent processing, establishing a causal link between NMDA receptors and recurrent processing in humans has remained challenging. Here, we report two pharmacological studies with randomized, double-blind, crossover designs in which we administered the NMDA antagonist memantine, while collecting human electroencephalography (EEG). We trained and tested EEG classifiers to reflect the processing of specific stimulus features with increasing levels of complexity, namely differences in stimulus contrast, collinearity between local line elements, and illusory surfaces of a Kanizsa triangle. In two experiments involving different participants and visual tasks, we found that memantine selectively improved decoding of the Kanizsa illusion, known to depend on recurrent processing, while leaving decoding of contrast and collinearity largely unaffected. Interestingly, the results from an attentional blink (experiment 1) and task-relevance manipulation (experiment 2) showed that memantine was only effective when the stimulus was attended and consciously accessed. These findings suggest that NMDA inhibition through memantine enhances recurrent processing, especially for attended objects, and thereby provide a crucial step toward bridging animal and human research, shedding light on the neural mechanisms underpinning perceptual inference and conscious perception. This research was supported by a grant from the H2020 European Research Council (ERC STG 715605, SVG).

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bioRxiv2024

Criterion placement threatens the construct validity of neural measures of consciousness

How consciousness arises from brain activity has been a topic of intense scientific research for decades. But how does one identify the neural basis of something that is intrinsically personal and subjective? A hallmark approach has been to ask observers to judge stimuli as seen (conscious) and unseen (unconscious) and use post hoc sorting of neural measurements based these judgments. Unfortunately, cognitive and response biases are known to strongly affect how observers place their criterion for judging stimuli as seen vs. unseen, thereby confounding neural measures of consciousness. Surprisingly however, the effect of conservative and liberal criterion placement on neural measures of unconscious and conscious processing has never been explicitly investigated. Here we use simulations and electrophysiological brain measurements to show that conservative criterion placement has an unintuitive consequence: rather than selectively providing a cautious estimate of conscious processing, it inflates effect sizes in neural measures of both conscious and unconscious processing, while liberal criterion placement does the reverse. After showing this in simulation, we performed decoding analyses on two electroencephalography studies that employ common subjective indicators of conscious awareness, in which we experimentally manipulated the response criterion. The results confirm that the predicted confounding effects of criterion placement on neural measures of unconscious and conscious processing occur in empirical data, while further showing that the most widely used subjective scale, the Perceptual Awareness Scale (PAS), does not guard against criterion confounds. Follow-up simulations explicate how the experimental context determines whether the relative confounding effect of criterion placement is larger in neural measures of either conscious or unconscious processing. We conclude that criterion placement threatens the construct validity of neural measures of conscious and unconscious processing.

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bioRxiv2023

Isolating feedforward, lateral, and feedback mechanisms underlying perceptual and attentional impairments of conscious access

This study investigates failures in conscious access resulting from either weak sensory input (perceptual impairments) or unattended input (attentional impairments). Participants viewed a Kanizsa stimulus with or without an illusory triangle within a rapid serial visual presentation of distractor stimuli. We designed a novel Kanizsa stimulus that contained additional ancillary features of different complexity (local contrast and collinearity) that were independently manipulated. Perceptual performance on the Kanizsa stimulus (presence vs. absence of an illusion) was equated between the perceptual (masking) and attentional (attentional blink) manipulation to circumvent common confounds related to conditional differences in task performance. We trained and tested classifiers on electroencephalogram (EEG) data to reflect the processing of specific stimulus features, with increasing levels of complexity. We show that late stages of processing ([~]200-250 ms), reflecting the integration of complex stimulus features (collinearity, illusory triangle), were impaired by masking but spared by the attentional blink. In contrast, decoding of local contrast (the spatial arrangement of stimulus features) was observed early in time ([~]80 ms) and was left largely unaffected by either manipulation. These results replicate previous work showing that feedforward processing is largely preserved under both perceptual and attentional impairments. Crucially however, under matched levels of performance, only attentional impairments left the processing of more complex visual features relatively intact, likely related to spared lateral and local feedback processes during inattention. These findings reveal distinct neural mechanisms associated with perceptual and attentional impairments and thus contribute to a comprehensive understanding of distinct neural stages leading to conscious access.

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bioRxiv2023

A disinhibitory circuit mechanism explains a general principle of peak performance during mid-level arousal

Perceptual decision-making is highly dependent on the momentary arousal state of the brain, which fluctuates over time on a scale of hours, minutes, and even seconds. The textbook relationship between momentary arousal and task performance is captured by an inverted U-shape, as put forward in the Yerkes-Dodson law (Yerkes and Dodson, 1908). This law suggests optimal performance at moderate levels of arousal, and impaired performance at low or high arousal levels. However, despite its popularity, the evidence for this relationship in humans is mixed at best. Here, we use pupil-indexed arousal and performance data from various perceptual decision-making tasks to provide converging evidence for the inverted U-shaped relationship between spontaneous arousal fluctuations and performance across different decision types (discrimination, detection) and sensory modalities (visual, auditory). To further understand this relationship, we built a neurobiologically plausible mechanistic model and show that it is possible to reproduce our findings by incorporating two types of interneurons that are both modulated by an arousal signal. The model architecture produces two dynamical regimes under the influence of arousal: one regime in which performance increases with arousal, and another regime in which performance decreases with arousal, together forming an inverted U-shaped arousal-performance relationship. We conclude that the inverted U-shaped arousal-performance relationship is a general and robust property of sensory processing. It might be brought about by the influence of arousal on two types of interneurons that together act as a disinhibitory pathway for the neural populations that encode the available sensory evidence used for the decision. Significance statementWhen people are repeatedly performing the exact same task, their performance fluctuates over time. One of the causes of this behavioral variability is spontaneous fluctuations in arousal. According to the Yerkes-Dodson law, task performance is optimal at moderate levels of species arousal, with impaired performance at very low or high arousal levels. However, until now, the evidence supporting this law as a general mechanism in human decision-making is mixed, and a neural mechanism that may explain the inverted U-shaped arousal-performance relationship is lacking. We show that the Yerkes-Dodson law is a general law that holds for human observers across decision-making tasks and settings. Furthermore, we present a simple and neurobiologically plausible mechanistic model that can explain its existence.

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bioRxiv2022

Catecholaminergic neuromodulation and selective attention jointly shape perceptual decision making

Perceptual decisions about sensory input are influenced by fluctuations in ongoing neural activity, most prominently driven by attention and neuromodulator systems. It is currently unknown if neuromodulator activity and attention differentially modulate perceptual decision-making and/or whether neuromodulatory systems in fact control attentional processes. To investigate the effects of two distinct neuromodulatory systems and spatial attention on perceptual decisions, we pharmacologically elevated cholinergic (through donepezil) and catecholaminergic (through atomoxetine) levels in humans performing a visuo-spatial attention task, while we measured electroencephalography (EEG). Both attention and catecholaminergic enhancement improved decision-making at the behavioral and algorithmic level, as reflected in increased perceptual sensitivity and the modulation of the drift rate parameter derived from drift diffusion modeling. Univariate analyses of EEG data time-locked to the attentional cue, the target stimulus, and the motor response, further revealed that attention and catecholaminergic enhancement both modulated pre-stimulus cortical excitability, cue- and stimulus-evoked sensory activity as well as parietal evidence accumulation signals. Interestingly, we observed both similar, unique, and interactive effects of attention and catecholaminergic neuromodulation on these behavioral, algorithmic, and neural markers of the decision-making process. Thereby, this study reveals an intricate relationship between attentional and catecholaminergic systems and advances our understanding about how these systems jointly shape various stages of perceptual decision-making.

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bioRxiv2020

Processing of subjectively and objectively invisible stimuli in human visual cortex

The study of unconscious processing requires a measure of conscious awareness. Awareness measures can be either subjective (based on participants report) or objective (based on perceptual performance). The preferred awareness measure depends on the theoretical position about consciousness, and may influence conclusions about the extent of unconscious processing and about the neural correlates of consciousness. We obtained fMRI measurements from 43 subjects while they viewed masked faces and houses that were either subjectively or objectively invisible. We show that neural representations of objectively invisible faces and houses are limited to visual (shape-related) object properties, while subjectively invisible stimuli are processed up to more abstract, categorical levels of representation. These results demonstrate that the hypothesized extent of unconscious information processing is determined by the measurement approach. Furthermore, our data show that subjective and objective approaches are associated with different neural correlates of consciousness and thus have implications for neural theories of consciousness.

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bioRxiv2020

Intact sensory processing but hampered conflict detection when stimulus input is task-irrelevant

Conflict detection in sensory input is central to adaptive human behavior. Perhaps unsurprisingly, past research has shown that conflict may even be detected in absence of conflict awareness, suggesting that conflict detection is an automatic process that does not require attention. To test the possibility of conflict processing in the absence of attention, we manipulated task relevance and response overlap of potentially conflicting stimulus features across six behavioral tasks. Multivariate analyses on human electroencephalographic data revealed neural signatures of conflict only when at least one feature of a conflicting stimulus was attended, regardless of whether that feature was part of the conflict, or overlaps with the response. In contrast, neural signatures of basic sensory processes were present even when a stimulus was completely unattended. These data reveal an attentional bottleneck at the level of objects, suggesting that object-based attention is a prerequisite for cognitive control operations involved in conflict detection.

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bioRxiv2020

A silent disco: Persistent entrainment of low-frequency neural oscillations underlies beat-based, but not memory-based temporal expectations

The brain uses temporal structure in the environment, like rhythm in music and speech, to predict the timing of events, thereby optimizing their processing and perception. Temporal expectations can be grounded in different aspects of the input structure, such as a regular beat or a predictable pattern. One influential account posits that a generic mechanism underlies beat-based and pattern-based expectations, namely entrainment of low frequency neural oscillations to rhythmic input, while other accounts assume different underlying neural mechanisms. Here, we addressed this outstanding issue by examining EEG activity and behavioral responses during silent periods following rhythmic auditory sequences. We measured responses outlasting the rhythms both to avoid confounding the EEG analyses with evoked responses, and to directly test whether beat-based and pattern-based expectations persist beyond stimulation, as predicted by entrainment theories. To properly disentangle beat-based and pattern-based expectations, which often occur simultaneously, we used non-isochronous rhythms with a beat, a predictable pattern, or random timing. In Experiment 1 (N = 32), beat-based expectations affected behavioral ratings of probe events for two beat-cycles after the end of the rhythm. The effects of pattern-based expectations reflected expectations for one interval. In Experiment 2 (N = 27), using EEG, we found enhanced spectral power at the beat frequency for beat-based sequences both during listening and silence. For pattern-based sequences, enhanced power at a pattern-specific frequency was present during listening, but not silence. Moreover, we found a difference in the evoked signal following pattern-based and beat-based sequences. Finally, we show how multivariate pattern decoding and multi scale entropy - measures sensitive to non-oscillatory components of the signal - can be used to probe temporal expectations. Together, our results suggest that the input structure used to form temporal expectations may affect the associated neural mechanisms. We suggest climbing activity and low-frequency oscillations may be differentially associated with pattern-based and beat-based expectations.

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bioRxiv2019

Boosting Brain Signal Variability Underlies Liberal Shifts in Decision Bias

Strategically adopting decision biases allows organisms to tailor their choices to environmental demands. For example, a liberal response strategy pays off when target detection is crucial, whereas a conservative strategy is optimal for avoiding false alarms. Using conventional time-frequency analysis of human electroencephalographic (EEG) activity, we previously showed that bias setting entails adjustment of evidence accumulation in sensory regions (Kloosterman et al., 2019), but the presumed prefrontal signature of a strategic conservative-to-liberal bias shift has remained elusive. Here, we show that a liberal bias shift relies on frontal regions adopting a more unconstrained neural regime (boosted entropy) that is suited to the detection of unpredictable events. Overall EEG variation, spectral power and event-related potentials could not explain this relationship, highlighting the unique contribution of moment-to-moment neural variability to bias shifts. Neural variability modulation through prefrontal cortex appears instrumental for permitting an organism to tailor its decision bias to environmental demands. Impact statementMoment-to-moment variability is a prominent feature of neural activity. Rather than representing mere noise, this variability might enable us to flexibly adapt our decision biases to the environment.

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bioRxiv2019

Decreased alertness reconfigures cognitive control networks

Humans remarkable capacity to flexibly adapt their behaviour based on rapid situational changes is termed cognitive control. Intuitively, cognitive control is thought to be affected by the state of alertness, for example, when drowsy we feel less capable of adequately implementing effortful cognitive tasks. Although scientific investigations have focused on the effects of sleep deprivation and circadian time, little is known about how natural daily fluctuations in alertness in the regular awake state affect cognitive control. Here we combined a conflict task in the auditory domain with EEG neurodynamics to test how neural and behavioural markers of conflict processing are affected by fluctuations in alertness. Using a novel computational method, we segregated alert and drowsy trials from two testing sessions and observed that, although participants (both sexes) were generally sluggish, the typical Conflict Effect reflected in slower responses to conflicting information compared to non-conflicting information was still intact, as well as the moderating effect of previous conflict (Conflict Adaptation). However, the typical neural markers of cognitive control-local midfrontal-theta band power changes-that participants show during full alertness were no longer noticeable when alertness decreased. Instead, when drowsy, we found an increase in long-range information sharing (connectivity) between brain regions in the same frequency band. These results show the resilience of the human cognitive control system when affected by internal fluctuations of alertness, and suggest neural compensatory mechanisms at play in response to physiological pressure during diminished alertness. Significance StatementThe normal variability in alertness we experience in daily tasks is rarely taking into account in cognitive neuroscience. Here we studied neurobehavioral dynamics of cognitive control with decreasing alertness. We used the classic Simon Task where participants hear the word "left" or "right" in the right or left ear, eliciting slower responses when the word and the side are incongruent - the conflict effect. Participants performed the task both while fully awake and while getting drowsy, allowing for the characterisation of alertness modulating cognitive control. The changes in the neural signatures of conflict from local theta oscillations to a long-distance distributed theta network suggests a reconfiguration of the underlying neural processes subserving cognitive control when affected by alertness fluctuations.

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bioRxiv2019

Broadband Signal Rather than Frequency-Specific Rhythms Underlies Prediction Error in the Primate Auditory Cortex

Detection of statistical irregularities, measured as a prediction error response, is fundamental to the perceptual monitoring of the environment. We studied whether prediction error response is associated with neural oscillations or asynchronous broadband activity. Electrocorticography (ECoG) was carried out in three male monkeys, who passively listened to the auditory roving oddball stimuli. Local field potentials (LFP) recorded over the auditory cortex underwent spectral principal component analysis, which decoupled broadband and rhythmic components of the LFP signal. We found that the broadband component captured the prediction error response, whereas none of the rhythmic components were associated with statistical irregularities of sounds. The broadband component displayed more stochastic, asymmetrical multifractal properties than the rhythmic components, which revealed more self-similar dynamics. We thus conclude that the prediction error response is captured by neuronal populations generating asynchronous broadband activity, defined by irregular dynamical states which, unlike oscillatory rhythms, appear to enable the neural representation of auditory prediction error response. Significance StatementThis study aimed to examine the contribution of oscillatory and asynchronous components of auditory local field potentials in the generation of prediction error responses to sensory irregularities, as this has not been directly addressed in the previous studies. Here, we show that mismatch negativity - an auditory prediction error response - is driven by the asynchronous broadband component of potentials recorded in the auditory cortex. This finding highlights the importance of non-oscillatory neural processes in the predictive monitoring of the environment. At a more general level, the study demonstrates that stochastic neural processes, which are often disregarded as neural noise, do have a functional role in the processing of sensory information.

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bioRxiv2019

Multivariate analysis of EEG activity indexes contingent and non-contingent attentional capture

It is well known that salient yet irrelevant singleton can capture attention, even when this is inconsistent with the current goals of the observer (Theeuwes, 1992; 2010). Others however have claimed that capture is critically contingent on the goals of the observer: Capture is strongly modulated (or even eliminated) when the irrelevant singleton does not match the target-defining properties (Folk, Remington, & Johnston, 1992). There has been a long-standing debate on whether attentional capture can be explained by goal-driven and/or stimulus-driven accounts. Here, we shed further light on this phenomenon by using EEG activity (raw EEG and alpha power) to provide a time-resolved index of attentional orienting. Participants searched for a target defined by a pre-specified color. The search display was preceded by a singleton cue that either matched the color of the upcoming target (contingent cues), or that appeared in an irrelevant color (non-contingent cues). Multivariate analysis of raw EEG and alpha power revealed preferential tuning to the location of both contingent and non-contingent cues, with a stronger bias towards contingent than non-contingent cues. The time course of these effects, however, depended on the neural signal. Raw EEG data revealed attentional orienting towards the cue early on in the trial (>156 ms), while alpha power revealed sustained spatial selection in the cued locations at a later moment in the trial (>250 ms). Moreover, while raw EEG showed stronger capture by contingent cues during this early time window, the advantage for contingent cues arose during a later time window in alpha band activity. Thus, our findings suggest that raw EEG activity and alpha-band power tap into distinct neural processes that index movements of covert spatial attention. Both signals provide clear neural evidence that both contingent and non-contingent cues can capture attention, and that this process is robustly shaped by the target-defining properties in the current block of trials.

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bioRxiv2019

Humans can efficiently look for, but not select multiple visual objects

The human brain recurrently prioritizes task-relevant over task-irrelevant visual information. A central, question is whether multiple objects can be prioritized simultaneously. To answer this, we let observers search for two colored targets among distractors. Crucially, we independently varied the number of target colors that observers anticipated, and the number of target colors actually used to distinguish the targets in the display. This enabled us to dissociate the preparation of selection mechanisms from the actual engagement of such mechanisms. Multivariate classification of electroencephalographic activity allowed us to track selection of each target separately across time. The results revealed only small neural and behavioral costs associated with preparing for selecting two objects, but substantial costs when engaging in selection. Further analyses suggest this cost is the consequence of neural competition resulting in limited parallel processing, rather than a serial bottleneck. The findings bridge diverging theoretical perspectives on capacity limitations of feature-based attention.

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bioRxiv2019

The Role of Proactive and Reactive Cognitive Control for Target Selection in Multiple-Target Search

Cognitive control can involve proactive (preparatory) and reactive (corrective) mechanisms. Using a gaze-contingent eye tracking paradigm combined with fMRI, we investigated the involvement of these different modes of control and their underlying neural networks, when switching between different targets in multiple-target search. Participants simultaneously searched for two possible targets presented among distractors, and selected one of them. In one condition, only one of the targets was available in each display, so that the choice was imposed, and reactive control would be required. In the other condition, both targets were present, giving observers free choice over target selection, and allowing for proactive control. Switch costs emerged only when targets were imposed and not when target selection was free. We found differential levels of activity in the frontoparietal control network depending on whether target switches were free or imposed. Furthermore, we observed core regions of the default mode network to be active during target repetitions, indicating reduced control on these trials. Free and imposed switches jointly activated parietal and posterior frontal cortices, while free switches additionally activated anterior frontal cortices. These findings highlight unique contributions of proactive and reactive control during visual search.

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bioRxiv2019

High-pass filtering artifacts in multivariate classification of neural time series data

0.BackgroundTraditionally, EEG/MEG data are high-pass filtered and baseline-corrected to remove slow drifts. Minor deleterious effects of high-pass filtering in traditional time-series analysis have been well-documented, including temporal displacements. However, its effects on time-resolved multivariate pattern classification analyses (MVPA) are largely unknown. New MethodTo prevent potential displacement effects, we extend an alternative method of removing slow drift noise - robust detrending - with a procedure in which we mask out all cortical events from each trial. We refer to this method as trial-masked robust detrending. ResultsIn both real and simulated EEG data of a working memory experiment, we show that both high-pass filtering and standard robust detrending create artifacts that result in the displacement of multivariate patterns into activity silent periods, particularly apparent in temporal generalization analyses, and especially in combination with baseline correction. We show that trial-masked robust detrending is free from such displacements. Comparison with Existing Method(s)Temporal displacement may emerge even with modest filter cut-off settings such as 0.05 Hz, and even in regular robust detrending. However, trial-masked robust detrending results in artifact-free decoding without displacements. Baseline correction may unwittingly obfuscate spurious decoding effects and displace them to the rest of the trial. ConclusionsDecoding analyses benefits from trial-masked robust detrending, without the unwanted side effects introduced by filtering or regular robust detrending. However, for sufficiently clean data sets and sufficiently strong signals, no filtering or detrending at all may work adequately. Implications for other types of data are discussed, followed by a number of recommendations.

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