Time for What? Dissociating Explicit Timing Tasks through Electrophysiological Signatures.

Estimating durations between hundreds of milliseconds and seconds is essential for several daily tasks. Explicit timing tasks, which require participants to estimate durations to make a comparison (time for perception) or to reproduce them (time for action), are often used to investigate psychological and neural timing mechanisms. Recent studies have proposed that mechanisms may depend on specific task requirements. In this study, we conducted electroencephalogram (EEG) recordings on human participants as they estimated intervals in different task contexts to investigate the extent to which timing mechanisms depend on the nature of the task. We compared the neural processing of identical visual reference stimuli in two different tasks, in which stimulus durations were either perceptually compared or motorically reproduced in separate experimental blocks. Using multivariate pattern analyses, we could successfully decode the duration and the task of reference stimuli. We found evidence for both overlapping timing mechanisms across tasks as well as recruitment of task-dependent processes for comparing intervals for different purposes. Our findings suggest both core and specialized timing functions are recruited to support explicit timing tasks.

Multimodal resting-state connectivity predicts affective neurofeedback performance.

Neurofeedback has been suggested as a potential complementary therapy to different psychiatric disorders. Of interest for this approach is the prediction of individual performance and outcomes. In this study, we applied functional connectivity-based modeling using electroencephalography (EEG) and functional near-infrared spectroscopy (fNIRS) modalities to (i) investigate whether resting-state connectivity predicts performance during an affective neurofeedback task and (ii) evaluate the extent to which predictive connectivity profiles are correlated across EEG and fNIRS techniques. The fNIRS oxyhemoglobin and deoxyhemoglobin concentrations and the EEG beta and gamma bands modulated by the alpha frequency band (beta-m-alpha and gamma-m-alpha, respectively) recorded over the frontal cortex of healthy subjects were used to estimate functional connectivity from each neuroimaging modality. For each connectivity matrix, relevant edges were selected in a leave-one-subject-out procedure, summed into "connectivity summary scores" (CSS), and submitted as inputs to a support vector regressor (SVR). Then, the performance of the left-out-subject was predicted using the trained SVR model. Linear relationships between the CSS across both modalities were evaluated using Pearson's correlation. The predictive model showed a mean absolute error smaller than 20%, and the fNIRS oxyhemoglobin CSS was significantly correlated with the EEG gamma-m-alpha CSS (r = -0.456, p = 0.030). These results support that pre-task electrophysiological and hemodynamic resting-state connectivity are potential predictors of neurofeedback performance and are meaningfully coupled. This investigation motivates the use of joint EEG-fNIRS connectivity as outcome predictors, as well as a tool for functional connectivity coupling investigation.

Time encoding migrates from prefrontal cortex to dorsal striatum during learning of a self-timed response duration task.

Although time is a fundamental dimension of life, we do not know how brain areas cooperate to keep track and process time intervals. Notably, analyses of neural activity during learning are rare, mainly because timing tasks usually require training over many days. We investigated how the time encoding evolves when animals learn to time a 1.5 s interval. We designed a novel training protocol where rats go from naive- to proficient-level timing performance within a single session, allowing us to investigate neuronal activity from very early learning stages. We used pharmacological experiments and machine-learning algorithms to evaluate the level of time encoding in the medial prefrontal cortex and the dorsal striatum. Our results show a double dissociation between the medial prefrontal cortex and the dorsal striatum during temporal learning, where the former commits to early learning stages while the latter engages as animals become proficient in the task.

Subject-independent decoding of affective states using functional near-infrared spectroscopy.

Affective decoding is the inference of human emotional states using brain signal measurements. This approach is crucial to develop new therapeutic approaches for psychiatric rehabilitation, such as affective neurofeedback protocols. To reduce the training duration and optimize the clinical outputs, an ideal clinical neurofeedback could be trained using data from an independent group of volunteers before being used by new patients. Here, we investigated if this subject-independent design of affective decoding can be achieved using functional near-infrared spectroscopy (fNIRS) signals from frontal and occipital areas. For this purpose, a linear discriminant analysis classifier was first trained in a dataset (49 participants, 24.65±3.23 years) and then tested in a completely independent one (20 participants, 24.00±3.92 years). Significant balanced accuracies between classes were found for positive vs. negative (64.50 ± 12.03%, p<0.01) and negative vs. neutral (68.25 ± 12.97%, p<0.01) affective states discrimination during a reactive block consisting in viewing affective-loaded images. For an active block, in which volunteers were instructed to recollect personal affective experiences, significant accuracy was found for positive vs. neutral affect classification (71.25 ± 18.02%, p<0.01). In this last case, only three fNIRS channels were enough to discriminate between neutral and positive affective states. Although more research is needed, for example focusing on better combinations of features and classifiers, our results highlight fNIRS as a possible technique for subject-independent affective decoding, reaching significant classification accuracies of emotional states using only a few but biologically relevant features.

Prestimulus alpha power is related to the strength of stimulus representation.

Spatial attention can modulate behavioural performance and is associated with several electrophysiological markers. In this study, we used multivariate pattern analysis in electrophysiology data to investigate the effects of covert spatial attention on the quality of stimulus processing and its underlying mechanisms. Our results show that covert spatial attention led to (i) an anticipatory alpha power desynchronization; (ii) enhanced stimuli identity information. Moreover, we found that alpha power fluctuations in anticipation of the relevant stimuli boosted and prolonged the coding of stimulus identity.

Temporal regularity and stimulus control in multiple fixed interval schedules.

In multiple fixed interval schedules of reinforcement, different time intervals are signaled by different environmental stimuli which acquire control over behavior. Previous work has shown that temporal performance is controlled not only by external stimuli but also by temporal aspects of the task, depending on the order in which the different intervals are trained - intermixed across trials or in blocks of several trials. The aim of this study was to further describe the training conditions under which the stimuli acquire control over temporal performance. We manipulated the number of consecutive trials of each fixed interval (FI) per training block (Experiment I) and the number of FIs trained (Experiment II). The results suggest that when trained in blocks of several consecutive trials of the same FI, temporal performance is controlled by temporal regularities across trials and not by the visual stimuli that signal the FIs. One possible account for those data is that the temporal cues overshadow the visual stimuli for the control of temporal performance. Similar results have also been observed with humans, which suggest that temporal regularity overcomes the stimuli in the control of behavior in temporal tasks across species.

A common representation of time across visual and auditory modalities.

Humans' and non-human animals' ability to process time on the scale of milliseconds and seconds is essential for adaptive behaviour. A central question of how brains keep track of time is how specific temporal information across different sensory modalities is. In the present study, we show that encoding of temporal intervals in auditory and visual modalities are qualitatively similar. Human participants were instructed to reproduce intervals in the range from 750 ms to 1500 ms marked by auditory or visual stimuli. Our behavioural results suggest that, although participants were more accurate in reproducing intervals marked by auditory stimuli, there was a strong correlation in performance between modalities. Using multivariate pattern analysis in scalp EEG, we show that activity during late periods of the intervals was similar within and between modalities. Critically, we show that a multivariate pattern classifier was able to accurately predict the elapsed interval, even when trained on an interval marked by a stimulus of a different sensory modality. Taken together, our results suggest that, while there are differences in the processing of intervals marked by auditory and visual stimuli, they also share a common neural representation.

The effect of conscious intention to act on the Bereitschaftspotential.

The current study investigated the effect of conscious intention to act on the Bereitschaftspotential. Situations in which the awareness of acting is minimally expressed were generated by asking 16 participants to press a button after performing a mental imagery task based on animal pictures (automatic condition). The affective responses induced by the pictures were controlled by selecting the animals according to different valences, threatening and neutral. The Bereitschaftspotential associated with the button presses was compared to the observed when similar movements were performed under the basic instructions of the self-paced movement paradigm (willed condition). Enhanced Bereitschaftspotential amplitudes were observed in the willed condition with respect to the automatic condition. This effect was manifested as a negative slope at medial frontocentral sites during the last 500 ms before movement onset. The valence of the pictures did not affect the motor preparatory potentials. The results suggest that significant part of the NS' subcomponent of the readiness potential is associated with the attention to-and, presumably, awareness of-intention to move, possibly reflecting cortical activation from supplementary motor areas. Secondarily, our findings supports that the feeling of threat does not influence the Bereitschaftspotential associated with automatic movements. Regarding methodological issues, the behavioural model of spontaneous voluntary movements proposed in automatic condition can benefit investigations on purely motor (or non-cognitive) subcomponents of the Bereitschaftspotential.

Predicting affective valence using cortical hemodynamic signals.

Ascribing affective valence to stimuli or mental states is a fundamental property of human experiences. Recent neuroimaging meta-analyses favor the workspace hypothesis for the neural underpinning of valence, in which both positive and negative values are encoded by overlapping networks but are associated with different patterns of activity. In the present study, we further explored this framework using functional near-infrared spectroscopy (fNIRS) in conjunction with multivariate analyses. We monitored the fronto-temporal and occipital hemodynamic activity of 49 participants during the viewing of affective images (passive condition) and during the imagination of affectively loaded states (active condition). Multivariate decoding techniques were applied to determine whether affective valence is encoded in the cortical areas assessed. Prediction accuracies of 89.90 ± 13.84% and 85.41 ± 14.43% were observed for positive versus neutral comparisons, and of 91.53 ± 13.04% and 81.54 ± 16.05% for negative versus neutral comparisons (passive/active conditions, respectively). Our results are consistent with previous studies using other neuroimaging modalities that support the affective workspace hypothesis and the notion that valence is instantiated by the same network, regardless of whether the affective experience is passively or actively elicited.

Functional near-infrared spectroscopy-based affective neurofeedback: feedback effect, illiteracy phenomena, and whole-connectivity profiles.

Background: Affective neurofeedback constitutes a suitable approach to control abnormal neural activities associated with psychiatric disorders and might consequently relief symptom severity. However, different aspects of neurofeedback remain unclear, such as its neural basis, the performance variation, the feedback effect, among others. Aim: First, we aimed to propose a functional near-infrared spectroscopy (fNIRS)-based affective neurofeedback based on the self-regulation of frontal and occipital networks. Second, we evaluated three different feedback approaches on performance: real, fixed, and random feedback. Third, we investigated different demographic, psychological, and physiological predictors of performance. Approach: Thirty-three healthy participants performed a task whereby an amorphous figure changed its shape according to the elicited affect (positive or neutral). During the task, the participants randomly received three different feedback approaches: real feedback, with no change of the classifier output; fixed feedback, keeping the feedback figure unmodified; and random feedback, where the classifier output was multiplied by an arbitrary value, causing a feedback different than expected by the subject. Then, we applied a multivariate comparison of the whole-connectivity profiles according to the affective states and feedback approaches, as well as during a pretask resting-state block, to predict performance. Results: Participants were able to control this feedback system with 70.00 % ± 24.43 % ( p < 0.01 ) of performance during the real feedback trials. No significant differences were found when comparing the average performances of the feedback approaches. However, the whole functional connectivity profiles presented significant Mahalanobis distances ( p ≪ 0.001 ) when comparing both affective states and all feedback approaches. Finally, task performance was positively correlated to the pretask resting-state whole functional connectivity ( r = 0.512 , p = 0.009 ). Conclusions: Our results suggest that fNIRS might be a feasible tool to develop a neurofeedback system based on the self-regulation of affective networks. This finding enables future investigations using an fNIRS-based affective neurofeedback in psychiatric populations. Furthermore, functional connectivity profiles proved to be a good predictor of performance and suggested an increased effort to maintain task control in the presence of feedback distractors.

Temporal Anticipation Based on Memory.

The fundamental role that our long-term memories play in guiding perception is increasingly recognized, but the functional and neural mechanisms are just beginning to be explored. Although experimental approaches are being developed to investigate the influence of long-term memories on perception, these remain mostly static and neglect their temporal and dynamic nature. Here, we show that our long-term memories can guide attention proactively and dynamically based on learned temporal associations. Across two experiments, we found that detection and discrimination of targets appearing within previously learned contexts are enhanced when the timing of target appearance matches the learned temporal contingency. Neural markers of temporal preparation revealed that the learned temporal associations trigger specific temporal predictions. Our findings emphasize the ecological role that memories play in predicting and preparing perception of anticipated events, calling for revision of the usual conceptualization of contextual associative memory as a reflective and retroactive function.

Dynamic representation of time in brain states.

The ability to process time on the scale of milliseconds and seconds is essential for behaviour. A growing number of studies have started to focus on brain dynamics as a mechanism for temporal encoding. Although there is growing evidence in favour of this view from computational and in vitro studies, there is still a lack of results from experiments in humans. We show that high-dimensional brain states revealed by multivariate pattern analysis of human EEG are correlated to temporal judgements. First, we show that, as participants estimate temporal intervals, the spatiotemporal dynamics of their brain activity are consistent across trials. Second, we present evidence that these dynamics exhibit properties of temporal perception, such as scale invariance. Lastly, we show that it is possible to predict temporal judgements based on brain states. These results show how scalp recordings can reveal the spatiotemporal dynamics of human brain activity related to temporal processing.

Lexical-retrieval and semantic memory in Parkinson's disease: The question of noun and verb dissociation.

The dissociation between the processing of verbs and nouns has been debated in light of the Embodied Cognition Theory (EC). The objective of this paper is to verify how action and verb processing deficits of PD patients are modulated by different tasks with different cognitive demands. Action and object lexical-semantic processing was evaluated in patients with Parkinson's Disease (PD) and cognitively healthy controls through three different tasks (verbal fluency, naming and semantic association). Compared to controls, PD patients presented worse performance in naming actions and in the two semantic association tasks (action/object). Action verbal fluency performance was significantly associated with PD severity whereas object semantic association deficits and noun verbal fluency scores were associated to lower scores in measures of global cognitive functioning. Our data suggest that semantic deficits are related to the type of cognitive processing and this is in the line with more flexible EC accounts.

Experts in offside decision making learn to compensate for their illusory perceptions.

In association football, the flash-lag effect appears to be a viable explanation for erroneous offside decision making. Due to this spatiotemporal illusion, assistant referees (ARs) perceive the player who receives the ball ahead of his real position. In this experiment, a laboratory decision-making task was used to demonstrate that international top-class ARs, compared with amateur soccer players, do not have superior perceptual sensitivity. They clearly modify their decision criterion according to the contextual needs and, therefore, show a higher response bias toward not responding to the stimulus, in particular in the most difficult situations. Thus, international ARs show evidence for response-level compensation, resulting in a specific cost (i.e., more misses), which clearly reflects the use of particular (cognitive) strategies. In summary, it appears that experts in offside decision making can be distinguished from novices more on the cognitive or decision-making level than on the perceptual level.

Bias and learning in temporal binding: intervals between actions and outcomes are compressed by prior bias.

It has consistently been shown that agents judge the intervals between their actions and outcomes as compressed in time, an effect named intentional binding. In the present work, we investigated whether this effect is result of prior bias volunteers have about the timing of the consequences of their actions, or if it is due to learning that occurs during the experimental session. Volunteers made temporal estimates of the interval between their action and target onset (Action conditions), or between two events (No-Action conditions). Our results show that temporal estimates become shorter throughout each experimental block in both conditions. Moreover, we found that observers judged intervals between action and outcomes as shorter even in very early trials of each block. To quantify the decrease of temporal judgments in experimental blocks, exponential functions were fitted to participants' temporal judgments. The fitted parameters suggest that observers had different prior biases as to intervals between events in which action was involved. These findings suggest that prior bias might play a more important role in this effect than calibration-type learning processes.

Temporal expectation enhances contrast sensitivity by phase entrainment of low-frequency oscillations in visual cortex.

Although it is increasingly accepted that temporal expectation can modulate early perceptual processing, the underlying neural computations remain unknown. In the present study, we combined a psychophysical paradigm with electrophysiological recordings to investigate the putative contribution of low-frequency oscillatory activity in mediating the modulation of visual perception by temporal expectation. Human participants judged the orientation of brief targets (visual Gabor patterns tilted clockwise or counterclockwise) embedded within temporally regular or irregular streams of noise-patches used as temporal cues. Psychophysical results indicated that temporal expectation enhanced the contrast sensitivity of visual targets. A diffusion model indicated that rhythmic temporal expectation modulated the signal-to-noise gain of visual processing. The concurrent electrophysiological data revealed that the phase of delta oscillations overlying human visual cortex (1-4 Hz) was predictive of the quality of target processing only in regular streams of events. Moreover, in the regular condition, the optimum phase of these perception-predictive oscillations occurred in anticipation of the expected events. Together, these results show a strong correspondence between psychophysical and neurophysiological data, suggesting that the phase entrainment of low-frequency oscillations to external sensory cues can serve as an important and flexible mechanism for enhancing sensory processing.

Attention restores discrete items to visual short-term memory.

When a memory is forgotten, is it lost forever? Our study shows that selective attention can restore forgotten items to visual short-term memory (VSTM). In our two experiments, all stimuli presented in a memory array were designed to be equally task relevant during encoding. During the retention interval, however, participants were sometimes given a cue predicting which of the memory items would be probed at the end of the delay. This shift in task relevance improved recall for that item. We found that this type of cuing improved recall for items that otherwise would have been irretrievable, providing critical evidence that attention can restore forgotten information to VSTM. Psychophysical modeling of memory performance has confirmed that restoration of information in VSTM increases the probability that the cued item is available for recall but does not improve the representational quality of the memory. We further suggest that attention can restore discrete items to VSTM.

Temporal expectation improves the quality of sensory information.

It is increasingly clear that we extract patterns of temporal regularity between events to optimize information processing. Whereas some of the mechanisms for facilitating action preparation and execution have been well documented, much less is understood about whether and how temporal expectations influence visual perception. We used a psychophysical paradigm and computational modeling to investigate the mechanisms by which temporal expectation can modulate visual perception. Visual targets appeared in a stream of noise-patches separated by a fixed (400 ms regular condition) or jittered (200/300/400/500/600 ms irregular condition) intervals. Targets were visual gratings tilted 45° clockwise or counter-clockwise, presented at one of seven contrast levels. Human observers were required to perform an orientation discrimination (i.e., left or right). Psychometric functions for contrast sensitivity fitted for the regular and irregular conditions indicated that temporal expectation modulates perceptual processing by enhancing the contrast sensitivity of visual targets. This increase in the signal strength was accompanied by a reduction in reaction times. A diffusion model indicated that rhythmic temporal expectation enhanced the signal-to-noise gain of the sensory evidence upon which decisions were made. These effects support the idea that temporal structure of external events can entrain the attentional focus and psychophysical data, optimizing the processing of relevant sensory information.

Endogenous modulation of low frequency oscillations by temporal expectations.

Recent studies have associated increasing temporal expectations with synchronization of higher frequency oscillations and suppression of lower frequencies. In this experiment, we explore a proposal that low-frequency oscillations provide a mechanism for regulating temporal expectations. We used a speeded Go/No-go task and manipulated temporal expectations by changing the probability of target presentation after certain intervals. Across two conditions, the temporal conditional probability of target events differed substantially at the first of three possible intervals. We found that reactions times differed significantly at this first interval across conditions, decreasing with higher temporal expectations. Interestingly, the power of theta activity (4-8 Hz), distributed over central midline sites, also differed significantly across conditions at this first interval. Furthermore, we found a transient coupling between theta phase and beta power after the first interval in the condition with high temporal expectation for targets at this time point. Our results suggest that the adjustments in theta power and the phase-power coupling between theta and beta contribute to a central mechanism for controlling neural excitability according to temporal expectations.

The relation between action, predictability and temporal contiguity in temporal binding.

Previous studies have documented a subjective temporal attraction between actions and their effects. This finding, named intentional binding, is thought to be the result of a cognitive function that links actions to their consequences. Although several studies have tried to outline the necessary and sufficient conditions for intentional binding, a quantitative comparison between the roles of temporal contiguity, predictability and voluntary action and the evaluation of their interactions is difficult due to the high variability of the temporal binding measurements. In the present study, we used a novel methodology to investigate the properties of intentional binding. Subjects judged whether an auditory stimulus, which could either be triggered by a voluntary finger lift or be presented after a visual temporal marker unrelated to any action, was presented synchronously with a reference stimulus. In three experiments, the predictability, the interval between action and consequence and the presence of action itself were manipulated. The results indicate that (1) action is a necessary condition for temporal binding; (2) a fixed interval between the two events is not sufficient to cause the effect and (3) only in the presence of voluntary action do temporal predictability and contiguity play a significant role in modulating the effect.These findings are discussed in the context of the relationship between intentional binding and temporal expectation.