Ewa Beldzik

DIURNAL PATTERNS OF ACTIVITY OF THE ORIENTING AND EXECUTIVE ATTENTION NEURONAL NETWORKS IN SUBJECTS PERFORMING A STROOP-LIKE TASK: A FUNCTIONAL MAGNETIC RESONANCE IMAGING STUDY

Attentional processes are fundamental to good cognitive functioning of human operators. The purpose of this study was to analyze the activity of neuronal networks involved in the orienting attention and executive control processes from the perspective of diurnal variability. Twenty-three healthy male volunteers meeting magnetic resonance (MR) inclusion criteria performed the Stroop Color-Word task (block design) in the MR scanner five times/day (06:00, 10:00, 14:00, 18:00, 22:00 h). The first scanning session was scheduled 1–1.5 h after waking. Between MR sessions, subjects performed simulated driving tasks in stable environmental conditions, with controlled physical activity and diet. Significant activation was found in brain regions related to the orienting attentional system: the parietal lobe (BA40) and frontal eye-fields (FEFs). There were also activations in areas of the executive control system: the fronto-insular cortex (FIC), dorsal anterior cingulate cortex (dACC), presupplementary motor area (preSMA), supplementary motor area (SMA), basal ganglia, middle temporal (MT; BA21), and dorsolateral prefrontal cortex (DLPFC), as a part of the central executive network. Significant deactivations were observed in the rostral anterior cingulate cortex (rACC), posterior cingulate cortex (PCC), superior frontal gyrus (SF), parietal lobe (BA39), and parahippocampal that are thought to comprise the default mode network (DMN). Additionally, the activated regions included bilaterally lingual gyrus and fusiform gyrus. The insula was bilaterally deactivated. Visual attention controlled by the goal-oriented attention system and comprising top-down and bottom-up mechanisms, activated by Stroop-like task, turned out to be prone to diurnal changes. The study results show the occurrence of time-of-day–related variations in neural activity of brain regions linked to the orienting attentional system (left parietal lobe—BA40, left and right FEFs), simultaneously providing arguments for temporal stability of the executive system and default mode network. These results also seem to suggest that the involuntary, exogenous (bottom-up) mechanism of attention is more vulnerable to circadian and fatigue factors than the voluntary (top-down) mechanism, which appear to be maintained at the same functional level during the day. The above phenomena were observed at the neural level. (Author correspondence: marek@uj.edu.pl)

Neural networks related to pro-saccades and anti-saccades revealed by independent component analysis

The saccadic eye movement system provides an excellent model for investigating basic cognitive processes and flexible control over behaviour. While the mechanism of pro-saccades (PS) is well known, in the case of the anti-saccade task (AS) it is still not clear which brain regions play a role in the inhibition of reflexive saccade to the target, nor what is the exact mechanism of vector inversion (i.e. orienting in the opposite direction). Independent component analysis (ICA) is one of the methods being used to establish temporally coherent brain regions, i.e. neural networks related to the task. In the present study ICA was applied to fMRI data from PS and AS experiments. The study revealed separate networks responsible for saccade generation into the desired direction, the inhibition of automatic responses, as well as vector inversion. The first function is accomplished by the eye fields network. The inhibition of automatic responses is associated with the executive control network. Vector inversion seems to be accomplished by the network comprising a large set of areas, including intraparietal sulcus, precuneus/posterior cingulate cortices, retrosplenial and parahippocampal. Those regions are associated with the parieto-medial temporal pathway, so far linked only to navigation. These results provide a new insight into understanding of the processes of the inhibition and vector inversion.

Respiration phase-locks to fast stimulus presentations: Implications for the interpretation of posterior midline “deactivations”

The posterior midline region (PMR)—considered a core of the default mode network—is deactivated during successful performance in different cognitive tasks. The extent of PMR‐deactivations is correlated with task‐demands and associated with successful performance in various cognitive domains. In the domain of episodic memory, functional MRI (fMRI) studies found that PMR‐deactivations reliably predict learning (successful encoding). Yet it is unclear what explains this relation. One intriguing possibility is that PMR‐deactivations are partially mediated by respiratory artifacts. There is evidence that the fMRI signal in PMR is particularly prone to respiratory artifacts, because of its large surrounding blood vessels. As respiratory fluctuations have been shown to track changes in attention, it is critical for the general interpretation of fMRI results to clarify the relation between respiratory fluctuations, cognitive performance, and fMRI signal. Here, we investigated this issue by measuring respiration during word encoding, together with a breath‐holding condition during fMRI‐scanning. Stimulus‐locked respiratory analyses showed that respiratory fluctuations predicted successful encoding via a respiratory phase‐locking mechanism. At the same time, the fMRI analyses showed that PMR‐deactivations associated with learning were reduced during breath‐holding and correlated with individual differences in the respiratory phase‐locking effect during normal breathing. A left frontal region—used as a control region—did not show these effects. These findings indicate that respiration is a critical factor in explaining the link between PMR‐deactivation and successful cognitive performance. Further research is necessary to demonstrate whether our findings are restricted to episodic memory encoding, or also extend to other cognitive domains. Hum Brain Mapp 35:4932–4943, 2014.

Scale-Free Fluctuations in Behavioral Performance: Delineating Changes in Spontaneous Behavior of Humans with Induced Sleep Deficiency

The timing and dynamics of many diverse behaviors of mammals, e.g., patterns of animal foraging or human communication in social networks exhibit complex self-similar properties reproducible over multiple time scales. In this paper, we analyze spontaneous locomotor activity of healthy individuals recorded in two different conditions: during a week of regular sleep and a week of chronic partial sleep deprivation. After separating activity from rest with a pre-defined activity threshold, we have detected distinct statistical features of duration times of these two states. The cumulative distributions of activity periods follow a stretched exponential shape, and remain similar for both control and sleep deprived individuals. In contrast, rest periods, which follow power-law statistics over two orders of magnitude, have significantly distinct distributions for these two groups and the difference emerges already after the first night of shortened sleep. We have found steeper distributions for sleep deprived individuals, which indicates fewer long rest periods and more turbulent behavior. This separation of power-law exponents is the main result of our investigations, and might constitute an objective measure demonstrating the severity of sleep deprivation and the effects of sleep disorders.

Different Types of Errors in Saccadic Task Are Sensitive to Either Time of Day or Chronic Sleep Restriction

Circadian rhythms and restricted sleep length affect cognitive functions and, consequently, the performance of day to day activities. To date, no more than a few studies have explored the consequences of these factors on oculomotor behaviour. We have implemented a spatial cuing paradigm in an eye tracking experiment conducted four times of the day after one week of rested wakefulness and after one week of chronic partial sleep restriction. Our aim was to verify whether these conditions affect the number of a variety of saccadic task errors. Interestingly, we found that failures in response selection, i.e. premature responses and direction errors, were prone to time of day variations, whereas failures in response execution, i.e. omissions and commissions, were considerably affected by sleep deprivation. The former can be linked to the cue facilitation mechanism, while the latter to wake state instability and the diminished ability of top-down inhibition. Together, these results may be interpreted in terms of distinctive sensitivity of orienting and alerting systems to fatigue. Saccadic eye movements proved to be a novel and effective measure with which to study the susceptibility of attentional systems to time factors, thus, this approach is recommended for future research.

I Don’t Want to Miss a Thing – Learning Dynamics and Effects of Feedback Type and Monetary Incentive in a Paired Associate Deterministic Learning Task

Effective functioning in a complex environment requires adjusting of behavior according to changing situational demands. To do so, organisms must learn new, more adaptive behaviors by extracting the necessary information from externally provided feedback. Not surprisingly, feedback-guided learning has been extensively studied using multiple research paradigms. The purpose of the present study was to test the newly designed Paired Associate Deterministic Learning task (PADL), in which participants were presented with either positive or negative deterministic feedback. Moreover, we manipulated the level of motivation in the learning process by comparing blocks with strictly cognitive, informative feedback to blocks where participants were additionally motivated by anticipated monetary reward or loss. Our results proved the PADL to be a useful tool not only for studying the learning process in a deterministic environment, but also, due to the varying task conditions, for assessing differences in learning patterns. Particularly, we show that the learning process itself is influenced by manipulating both the type of feedback information and the motivational significance associated with the expected monetary reward.

Seeking a fingerprint: analysis of point processes in actigraphy recording

Motor activity of humans displays complex temporal fluctuations which can be characterised by scale-invariant statistics, thus demonstrating that structure and fluctuations of such kinetics remain similar over a broad range of time scales. Previous studies on humans regularly deprived of sleep or suffering from sleep disorders predicted a change in the invariant scale parameters with respect to those for healthy subjects. In this study we investigate the signal patterns from actigraphy recordings by means of characteristic measures of fractional point processes. We analyse spontaneous locomotor activity of healthy individuals recorded during a week of regular sleep and a week of chronic partial sleep deprivation. Behavioural symptoms of lack of sleep can be evaluated by analysing statistics of duration times during active and resting states, and alteration of behavioural organisation can be assessed by analysis of power laws detected in the event count distribution, distribution of waiting times between consecutive movements and detrended fluctuation analysis of recorded time series. We claim that among different measures characterising complexity of the actigraphy recordings and their variations implied by chronic sleep distress, the exponents characterising slopes of survival functions in resting states are the most effective biomarkers distinguishing between healthy and sleep-deprived groups.

Brain Activations Related to Saccadic Response Conflict are not Sensitive to Time on Task

Establishing a role of the dorsal medial frontal cortex in the performance monitoring and cognitive control has been a challenge to neuroscientists for the past decade. In light of recent findings, the conflict monitoring hypothesis has been elaborated to an action-outcome predictor theory. One of the findings that led to this re-evaluation was the fMRI study in which conflict-related brain activity was investigated in terms of the so-called time on task effect, i.e., a linear increase of the BOLD signal with longer response times. The aim of this study was to investigate brain regions involved in the processing of saccadic response conflict and to account for the time on task effect. A modified spatial cueing task was implemented in the event-related fMRI study with oculomotor responses. The results revealed several brain regions which show higher activity for incongruent trials in comparison to the congruent ones, including pre-supplementary motor area together with the frontal and parietal regions. Further analysis accounting for the effect of response time provided evidence that these brain activations were not sensitive to time on task but reflected purely the congruency effect.

Dynamics of error-related activity in deterministic learning - an EEG and fMRI study

There is a close relationship between progress in learning and the error-monitoring system. EEG and fMRI studies using probabilistic learning have revealed the distinct dynamics of this system after response and feedback, i.e. an increase of error-related and a decrease of feedback-related activity in the anterior cingulate cortex (ACC). Based on the existing theories, it can be presumed that during deterministic learning feedback-related activity in ACC would also increase. Since these assumptions have not yet been confirmed, it can be only speculated based on the data from the probabilistic paradigms how the information is being integrated within the ACC during deterministic learning. Here we implemented the Paired Associate Deterministic Learning task to the EEG and fMRI experiments. The analysis of EEG data showed a significant increase in the amplitude for both ERN and FN. Similarly, the fMRI results showed an increase in response-related and feedback-related activity of the ACC in erroneous trials. Our findings are in line with the current theories of ACC function: increasing ACC activity can be linked to the detected discrepancy between expected and obtained outcomes. We argue that expectancy violations in the course of deterministic learning are signalled by both, internal and external evaluation system.

When three is greater than five : EEG and fMRI signatures of errors in numerical and physical comparisons

Unravelling the neural mechanisms, which determine performance accuracy, is one of the key concepts in cognitive neuroscience. When compared to correct responses, shorter reaction times are commonly observed behavioural feature of errors committed in typical conflict tasks. Yet, little is known about the origins of this phenomenon. In this study, EEG and fMRI experiments were conducted using the numerical version of the Stroop paradigm, which yielded unique behavioural outcomes. Particularly, errors in numerical comparison had shorter reaction times than correct trials, whereas physical comparison resulted in the opposite pattern. This criss–crossing interaction effect was used as a marker when exploring time-courses of brain activity. Group independent component analysis was applied to neurophysiological data and event-related analysis was conducted on the components’ time-courses. Results revealed one centro-parietal EEG component and one temporo-parietal fMRI neural network, which exhibited significant task and accuracy interactions. Showing linear increase that peaked right after the response onset, the activity of centro-parietal EEG component was linked to the decision variable signal, which reflects a process of accumulating evidence until reaching an action-triggering threshold. Both amplitude measurements and linear fits to the signal provided evidence for distinctive characteristics between numerical and physical comparisons, thereby explaining the behavioural outcomes: errors are committed due to accumulation of evidence in favour of the other (wrong) task instruction. The architecture of the temporo-parietal network, which comprises bilateral inferior temporal and intraparietal regions, is highly consistent with the recently established core “number network”. These findings link perceptual decisions with the generalized magnitude system and impart novel insights into the neural determinants of errors in humans.