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Kloosterman, N. A.

Publications and source records attributed to Kloosterman, N. A..

8 recordsLinked to original sources

Catecholamines reduce choice history biases

Theoretical accounts postulate that the catecholaminergic neuromodulator noradrenaline shapes cognition and behavior by reducing the impact of prior expectations on learning, inference, and decision-making. A ubiquitous effect of dynamic priors on perceptual decisions under uncertainty is choice history bias: the tendency to systematically repeat, or alternate, previous choices, even when stimulus categories are presented in a random sequence. Here, we directly test for a causal impact of catecholamines on these priors. We pharmacologically elevated catecholamine levels in human participants through the application of the noradrenaline reuptake inhibitor atomoxetine. We quantified the resulting changes in observers history biases in a visual perceptual decision task. Choice history biases in this task were highly idiosyncratic, tending toward choice repetition or alternation in different individuals. Atomoxetine decreased these biases (toward either repetition or alternation) compared to placebo. Behavioral modeling indicates that this bias reduction was due to a reduced bias in the accumulation of sensory evidence, rather than of the starting point of the accumulation process. Atomoxetine had no significant effect on other behavioral measures tested, including response time and choice accuracy. Atomoxetine and variations of pupil-linked arousal at slower and faster timescales had analogous effects on choice history bias. We conclude that catecholamines reduce the impact of a specific form of prior on perceptual decisions.

neuroscience↗

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.

neuroscience↗

Dynamic regulation of neural variability during working memory reflects dopamine, functional integration, and decision-making

The regulation of moment-to-moment neural variability may permit effective responses to changing cognitive demands. However, the mechanisms that support variability regulation are unknown. In the context of working memory, we leverage the largest available PET and fMRI dataset to jointly consider three lenses through which neural variability regulation could be understood: dopamine capacity, network-level functional integration, and flexible decision processes. We show that with greater working memory load, upregulation of variability was associated with elevated dopamine capacity and heightened functional integration, effects dominantly expressed in the striato-thalamic system rather than cortex. Strikingly, behavioral modeling revealed that working memory load evoked substantial decision biases during evidence accumulation, and those who jointly expressed a more optimal decision bias and higher dopamine capacity were most likely to upregulate striato-thalamic variability under load. We argue that the ability to align striato-thalamic variability to level of demand may be a hallmark of a well-functioning brain.

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Zapline-plus: a Zapline extension for automatic and adaptive removal of frequency-specific noise artifacts in M/EEG

Removing power line noise and other frequency-specific artifacts from electrophysiological data without affecting neural signals remains a challenging task. Recently, an approach was introduced that combines spectral and spatial filtering to effectively remove line noise: Zapline (de Cheveigne, 2020). This algorithm, however, requires manual selection of the noise frequency and the number of spatial components to remove during spatial filtering. Moreover, it assumes that noise frequency and spatial topography are stable over time, which is often not warranted. To overcome these issues, we introduce Zapline-plus, which allows adaptive and automatic removal of frequency-specific noise artifacts from M/EEG and LFP data. To achieve this, our extension first segments the data into periods (chunks) in which the noise is spatially stable. Then, for each chunk, it searches for peaks in the power spectrum, and finally applies Zapline. The exact noise frequency around the found target frequency is also determined separately for every chunk to allow fluctuations of the peak noise frequency over time. The number of to-be-removed components by Zapline is automatically determined using an outlier detection algorithm. Finally, the frequency spectrum after cleaning is analyzed for suboptimal cleaning and parameters are adapted accordingly if necessary before re-running the process. The software creates a detailed plot for monitoring the cleaning. We showcase the efficacy of the different features of our algorithm by applying it to four openly available data sets, two EEG sets containing both stationary and mobile task conditions, and two MEG sets containing strong line noise.

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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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Standard multiscale entropy reflects spectral power at mismatched temporal scales: What’s signal irregularity got to do with it?

Multiscale Entropy (MSE) is used to characterize the temporal irregularity of neural time series patterns. Due to its presumed sensitivity to non-linear signal characteristics, MSE is typically considered a complementary measure of brain dynamics to signal variance and spectral power. However, the divergence between these measures is often unclear in application. Furthermore, it is commonly assumed (yet sparingly verified) that entropy estimated at specific time scales reflects signal irregularity at those precise time scales of brain function. We argue that such assumptions are not tenable. Using simulated and empirical electroencephalogram (EEG) data from 47 younger and 52 older adults, we indicate strong and previously underappreciated associations between MSE and spectral power, and highlight how these links preclude traditional interpretations of MSE time scales. Specifically, we show that the typical definition of temporal patterns via "similarity bounds" biases coarse MSE scales - that are thought to reflect slow dynamics - by high-frequency dynamics. Moreover, we demonstrate that entropy at fine time scales - presumed to indicate fast dynamics - is highly sensitive to broadband spectral power, a measure dominated by low-frequency contributions. Jointly, these issues produce counterintuitive reflections of frequency-specific content on MSE time scales. We emphasize the resulting inferential problems in a conceptual replication of cross-sectional age differences at rest, in which scale-specific entropy age effects could be explained by spectral power differences at mismatched temporal scales. Furthermore, we demonstrate how such problems may be alleviated, resulting in the indication of scale-specific age differences in rhythmic irregularity. By controlling for narrowband contributions, we indicate that spontaneous alpha rhythms during eyes open rest transiently reduce broadband signal irregularity. Finally, we recommend best practices that may better permit a valid estimation and interpretation of neural signal irregularity at time scales of interest. Author SummaryBrain signals exhibit a wealth of dynamic patterns that that are thought to reflect ongoing neural computations. Multiscale sample entropy (MSE) intends to describe the temporal irregularity of such patterns at multiple time scales of brain function. However, the notion of time scales may often be unintuitive. In particular, traditional implementations of MSE are sensitive to slow fluctuations at fine time scales, and fast dynamics at coarse time scales. This conceptual divergence is often overlooked and may lead to difficulties in establishing the unique contribution of MSE to effects of interest over more established spectral power. Using simulations and empirical data, we highlight these issues and provide evidence for their relevance for valid practical inferences. We further highlight that standard MSE and traditional spectral power are highly collinear in our example. Finally, our analyses indicate that spectral filtering can be used to estimate temporal signal irregularity at matching and intuitive time scales. To guide future studies, we make multiple recommendations based on our observations. We believe that following these suggestions may advance our understanding of the unique contributions of neural signal irregularity to neural and cognitive function across the lifespan.

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Surprise About Sensory Event Timing Drives Cortical Transients in the Beta Frequency Band

Understanding the statistical structure of the environment is crucial for adaptive behavior. Humans and non-human decision-makers seem to track such structure through a process of probabilistic inference, which enables predictions about behaviorally relevant events. Deviations from such predictions cause surprise, which in turn helps improve the inference. Surprise about the timing of behaviorally relevant sensory events drives phasic responses of neuromodulatory brainstem systems, which project to the cerebral cortex. Here, we developed a computational model-based magnetoencephalography (MEG) approach for mapping the resulting cortical transients across space, time, and frequency, in the human brain. We used a Bayesian updating model to estimate the predicted timing of the next stimulus change in a simple visual detection task. This model yielded quantitative trial-by-trial estimates of temporal surprise. The model-based surprise variable predicted trial-by-trial variations in reaction time more strongly than the externally observable interval timings alone. Trial-by-trial variations in surprise were negatively correlated with the power of cortical population activity measured with MEG. This surprise-related power suppression occurred transiently around the behavioral response, specifically in the beta frequency band. It peaked in left lateral prefrontal as well as in frontal midline regions, and its cortical distribution was distinct from the movement-related suppression of beta power in motor cortex. Our results indicate that surprise about sensory event timing transiently suppresses ongoing beta-band oscillations in association cortex. This transient suppression of frontal beta-band oscillations might reflect an active reset triggered by surprise, and is in line with the idea that beta-oscillations help maintain cognitive sets.\n\nSignificance statementAgents continuously track the statistical structure of the environment, in order to make predictions about behaviorally relevant sensory events. Deviations from such predictions cause surprise, which in turn drives phasic responses of neuromodulatory brainstem systems that project to the cerebral cortex. We developed a computational model-based magnetoencephalography approach, which enabled us to map out transients changes in cortical population dynamics elicited by surprise about sensory event timing, across space, time, and frequency, in the human brain. The model-based estimates of surprise predicted behavior as well as a transient suppression of beta frequency-band oscillations in frontal cortical regions. Our results are in line with conceptual accounts that have linked neural oscillations in the beta-band to the maintenance of cognitive sets.

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Criterion Setting Modulates Neural Excitability of Human Visual Cortex

Decision bias is traditionally conceptualized as an internal reference against which sensory evidence is compared. Instead, we show that individuals implement decision bias by shifting the rate of sensory evidence accumulation towards a decision bound. Participants performed a target detection task while we recorded EEG. We experimentally manipulated participants decision criterion for reporting targets using different stimulus-response reward contingencies, inducing either a liberal or a conservative bias. Drift diffusion modeling revealed that a liberal strategy biased sensory evidence accumulation towards target-present choices. Moreover, a liberal bias resulted in stronger midfrontal pre-stimulus 2-6 Hz (theta) power and suppression of pre-stimulus 8--12 Hz (alpha) power in posterior cortex. The alpha suppression in turn mediated the output activity of visual cortex, as expressed in 59--100 Hz (gamma) power. These findings show that observers can intentionally control cortical excitability to strategically bias evidence accumulation towards the decision bound that maximizes their reward.

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