Human brain dynamics dissociate early perceptual and late motor-related stages of affordance processing.

Affordances, the opportunities for action offered by the environment to an agent, are vital for meaningful behaviour and exist in every interaction with the environment. There is an ongoing debate in the field about whether the perception of affordances is an automated process. Some studies suggest that affordance perception is an automated process that is independent from the visual context and bodily interaction with the environment, whereas others argue that it is modulated by the visual and motor context in which affordances are perceived. The present paper aims to resolve this debate by examining affordance automaticity from the perspective of sensorimotor time windows. To investigate the impact of different forms of bodily interactions with an environment, that is, the movement context (physical vs. joystick movement), we replicated a previous study on affordance perception in which participants actively moved through differently wide doors in an immersive 3D virtual environment. In the present study, we displayed the same environment on a 2D screen with participants moving through doors of different widths using the keys on a standard keyboard. We compared components of the event-related potential (ERP) from the continuously recorded electroencephalogram (EEG) that were previously reported to be related to affordance perception of architectural transitions (passable and impassable doors). Comparing early sensory and later motor-related ERPs, our study replicated ERPs reflecting early affordance perception but found differences in later motor-related components. These results indicate a shift from automated perception of affordances during early sensorimotor time windows to movement context dependence of affordance perception at later stages, suggesting that affordance perception is a dynamic and flexible process that changes over sensorimotor stages.

Sensorimotor brain dynamics reflect architectural affordances.

Anticipating meaningful actions in the environment is an essential function of the brain. Such predictive mechanisms originate from the motor system and allow for inferring actions from environmental affordances, and the potential to act within a specific environment. Using architecture, we provide a unique perspective on the ongoing debate in cognitive neuroscience and philosophy on whether cognition depends on movement or is decoupled from our physical structure. To investigate cognitive processes associated with architectural affordances, we used a mobile brain/body imaging approach recording brain activity synchronized to head-mounted displays. Participants perceived and acted on virtual transitions ranging from nonpassable to easily passable. We found that early sensory brain activity, on revealing the environment and before actual movement, differed as a function of affordances. In addition, movement through transitions was preceded by a motor-related negative component that also depended on affordances. Our results suggest that potential actions afforded by an environment influence perception.

The impact of hand movement velocity on cognitive conflict processing in a 3D object selection task in virtual reality.

Detecting and correcting incorrect body movements is an essential part of everyday interaction with one's environment. The human brain provides a monitoring system that constantly controls and adjusts our actions according to our surroundings. However, when our brain's predictions about a planned action do not match the sensory inputs resulting from that action, cognitive conflict occurs. Much is known about cognitive conflict in 1D/2D environments; however, less is known about the role of movement characteristics associated with cognitive conflict in 3D environment. Hence, we devised an object selection task in a virtual reality (VR) environment to test how the velocity of hand movements impacts human brain responses. From a series of analyses of EEG recordings synchronized with motion capture, we found that the velocity of the participants' hand movements modulated the brain's response to proprioceptive feedback during the task and induced a prediction error negativity (PEN). Additionally, the PEN originates in the anterior cingulate cortex and is itself modulated by the ballistic phase of the hand's movement. These findings suggest that velocity is an essential component of integrating hand movements with visual and proprioceptive information during interactions with real and virtual objects.

Single-trial regression of spatial exploration behavior indicates posterior EEG alpha modulation to reflect egocentric coding.

Learning to navigate uncharted terrain is a key cognitive ability that emerges as a deeply embodied process, with eye movements and locomotion proving most useful to sample the environment. We studied healthy human participants during active spatial learning of room-scale virtual reality (VR) mazes. In the invisible maze task, participants wearing a wireless electroencephalography (EEG) headset were free to explore their surroundings, only given the objective to build and foster a mental spatial representation of their environment. Spatial uncertainty was resolved by touching otherwise invisible walls that were briefly rendered visible inside VR, similar to finding your way in the dark. We showcase the capabilities of mobile brain/body imaging using VR, demonstrating several analysis approaches based on general linear models (GLMs) to reveal behavior-dependent brain dynamics. Confirming spatial learning via drawn sketch maps, we employed motion capture to image spatial exploration behavior describing a shift from initial exploration to subsequent exploitation of the mental representation. Using independent component analysis, the current work specifically targeted oscillations in response to wall touches reflecting isolated spatial learning events arising in deep posterior EEG sources located in the retrosplenial complex. Single-trial regression identified significant modulation of alpha oscillations by the immediate, egocentric, exploration behavior. When encountering novel walls, as well as with increasing walking distance between subsequent touches when encountering novel walls, alpha power decreased. We conclude that these oscillations play a prominent role during egocentric evidencing of allocentric spatial hypotheses.

Identifying key factors for improving ICA-based decomposition of EEG data in mobile and stationary experiments.

Recent developments in EEG hardware and analyses approaches allow for recordings in both stationary and mobile settings. Irrespective of the experimental setting, EEG recordings are contaminated with noise that has to be removed before the data can be functionally interpreted. Independent component analysis (ICA) is a commonly used tool to remove artifacts such as eye movement, muscle activity, and external noise from the data and to analyze activity on the level of EEG effective brain sources. The effectiveness of filtering the data is one key preprocessing step to improve the decomposition that has been investigated previously. However, no study thus far compared the different requirements of mobile and stationary experiments regarding the preprocessing for ICA decomposition. We thus evaluated how movement in EEG experiments, the number of channels, and the high-pass filter cutoff during preprocessing influence the ICA decomposition. We found that for commonly used settings (stationary experiment, 64 channels, 0.5 Hz filter), the ICA results are acceptable. However, high-pass filters of up to 2 Hz cut-off frequency should be used in mobile experiments, and more channels require a higher filter to reach an optimal decomposition. Fewer brain ICs were found in mobile experiments, but cleaning the data with ICA has been proved to be important and functional even with low-density channel setups. Based on the results, we provide guidelines for different experimental settings that improve the ICA decomposition.

Eye movement-related brain potentials during assisted navigation in real-world environments.

Conducting neuroscience research in the real-world remains challenging because of movement- and environment-related artifacts as well as missing control over stimulus presentation. The present study overcame these restrictions by mobile electroencephalography (EEG) and data-driven analysis approaches during a real-world navigation task. During assisted navigation through an unfamiliar city environment, participants received either standard or landmark-based auditory navigation instructions. EEG data were recorded continuously during navigation. Saccade- and blink-events as well as gait-related EEG activity were extracted from sensor level data. Brain activity associated with the navigation task was identified by subsequent source-based cleaning of non-brain activity and unfolding of overlapping event-related potentials. When navigators received landmark-based instructions compared to those receiving standard navigation instructions, the blink-related brain potentials during navigation revealed higher amplitudes at fronto-central leads in a time window starting at 300 ms after blinks, which was accompanied by improved spatial knowledge acquisition tested in follow-up spatial tasks. Replicating improved spatial knowledge acquisition from previous experiments, the present study revealed eye movement-related brain potentials to point to the involvement of higher cognitive processes and increased processing of incoming information during periods of landmark-based instructions. The study revealed neuronal correlates underlying visuospatial information processing during assisted navigation in the real-world providing a new analysis approach for neuroscientific research in freely moving participants in uncontrollable real-world environments.

The AudioMaze: An EEG and motion capture study of human spatial navigation in sparse augmented reality.

Spatial navigation is one of the fundamental cognitive functions central to survival in most animals. Studies in humans investigating the neural foundations of spatial navigation traditionally use stationary, desk-top protocols revealing the hippocampus, parahippocampal place area (PPA), and retrosplenial complex to be involved in navigation. However, brain dynamics, while freely navigating the real world remain poorly understood. To address this issue, we developed a novel paradigm, the AudioMaze, in which participants freely explore a room-sized virtual maze, while EEG is recorded synchronized to motion capture. Participants (n = 16) were blindfolded and explored different mazes, each in three successive trials, using their right hand as a probe to "feel" for virtual maze walls. When their hand "neared" a virtual wall, they received directional noise feedback. Evidence for spatial learning include shortening of time spent and an increase of movement velocity as the same maze was repeatedly explored. Theta-band EEG power in or near the right lingual gyrus, the posterior portion of the PPA, decreased across trials, potentially reflecting the spatial learning. Effective connectivity analysis revealed directed information flow from the lingual gyrus to the midcingulate cortex, which may indicate an updating process that integrates spatial information with future action. To conclude, we found behavioral evidence of navigational learning in a sparse-AR environment, and a neural correlate of navigational learning was found near the lingual gyrus.

Alteration of brain dynamics during dual-task overground walking.

When walking in our natural environment, we often solve additional cognitive tasks. This increases the demand of resources needed for both the cognitive and motor systems, resulting in Cognitive-Motor Interference (CMI). A large portion of neurophysiological investigations on CMI took place in static settings, emphasizing the experimental rigor but overshadowing the ecological validity. As a more ecologically valid alternative to treadmill and desktop-based setups to investigate CMI, we developed a dual-task walking scenario in virtual reality (VR) combined with Mobile Brain/Body Imaging (MoBI). We aimed at investigating how brain dynamics are modulated by dual-task overground walking with an additional task in the visual domain. Participants performed a visual discrimination task in VR while standing (single-task) and walking overground (dual-task). Even though walking had no impact on the performance in the visual discrimination task, a P3 amplitude reduction along with changes in power spectral densities (PSDs) were observed for discriminating visual stimuli during dual-task walking. These results reflect an impact of walking on the parallel processing of visual stimuli even when the cognitive task is particularly easy. This standardized and easy to modify VR paradigm helps to systematically study CMI, allowing researchers to control for the impact of additional task complexity of tasks in different sensory modalities. Future investigations implementing an improved virtual design with more challenging cognitive and motor tasks will have to investigate the roles of both cognition and motion, allowing for a better understanding of the functional architecture of attention reallocation between cognitive and motor systems during active behavior.

Mobile brain/body Imaging in dance: A dynamic transdisciplinary field for applied research.

Neuroscience of dance is an emerging field with important applications related to health and well-being, as dance has shown potential to foster adaptive neuroplasticity and is increasingly popular as a therapeutic activity or adjunct therapy for people living with conditions such as Parkinson's and Alzheimer's diseases. However, the multimodal nature of dance presents challenges to researchers aiming to identify mechanisms involved when dance is used to combat neurodegeneration or support healthy ageing. Requiring simultaneous engagement of motor and cognitive domains, dancing includes coordination of systems involved in timing, memory and spatial learning. Studies on dance to this point rely primarily on assessments of brain dynamics and structure through pre/post-tests or studies on expertise, as traditional brain imaging modalities restrict participant movement to avoid movement-related artefacts. In this paper, we describe the process of designing and implementing a study that uses mobile brain/body imaging (MoBI) to investigate real-time changes in brain dynamics and behaviour during the process of learning and performing a novel dance choreography. We show the potential for new insights to emerge from the coordinated collection of movement and brain-based data, and the implications of these in an emerging field whose medium is motion.

Time to move: Brain dynamics underlying natural action and cognition.

Advances in Mobile Brain/Body Imaging (MoBI) technology allows for real-time measurements of human brain dynamics during every day, natural, real-life situations. This special issue Time to Move brings together a collection of experimental papers, targeted reviews and opinion articles that lay out the latest MoBI findings. A wide range of topics across different fields are covered including art, athletics, virtual reality, and mobility. What unites these diverse topics is the common goal to enhance and restore human abilities by reaching a better understanding on how cognition is implemented by the brain-body relationship. The breadth and novelty of paradigms and findings reported here positions MoBI as a new frontier in the field of human cognitive neuroscience.

Mobile brain/body imaging of landmark-based navigation with high-density EEG.

Coupling behavioral measures and brain imaging in naturalistic, ecological conditions is key to comprehend the neural bases of spatial navigation. This highly integrative function encompasses sensorimotor, cognitive, and executive processes that jointly mediate active exploration and spatial learning. However, most neuroimaging approaches in humans are based on static, motion-constrained paradigms and they do not account for all these processes, in particular multisensory integration. Following the Mobile Brain/Body Imaging approach, we aimed to explore the cortical correlates of landmark-based navigation in actively behaving young adults, solving a Y-maze task in immersive virtual reality. EEG analysis identified a set of brain areas matching state-of-the-art brain imaging literature of landmark-based navigation. Spatial behavior in mobile conditions additionally involved sensorimotor areas related to motor execution and proprioception usually overlooked in static fMRI paradigms. Expectedly, we located a cortical source in or near the posterior cingulate, in line with the engagement of the retrosplenial complex in spatial reorientation. Consistent with its role in visuo-spatial processing and coding, we observed an alpha-power desynchronization while participants gathered visual information. We also hypothesized behavior-dependent modulations of the cortical signal during navigation. Despite finding few differences between the encoding and retrieval phases of the task, we identified transient time-frequency patterns attributed, for instance, to attentional demand, as reflected in the alpha/gamma range, or memory workload in the delta/theta range. We confirmed that combining mobile high-density EEG and biometric measures can help unravel the brain structures and the neural modulations subtending ecological landmark-based navigation.

Validation of a Low-Cost Electrocardiography (ECG) System for Psychophysiological Research.

BACKGROUND AND OBJECTIVE: The reliability of low-cost mobile systems for recording Electrocardiographic (ECG) data is mostly unknown, posing questions regarding the quality of the recorded data and the validity of the extracted physiological parameters. The present study compared the BITalino toolkit with an established medical-grade ECG system (BrainAmp-ExG). METHODS: Participants underwent simultaneous ECG recordings with the two instruments while watching pleasant and unpleasant pictures of the "International Affective Picture System" (IAPS). Common ECG parameters were extracted and compared between the two systems. The Intraclass Correlation Coefficients (ICCs) and the Bland-Altman Limits of Agreement (LoA) method served as criteria for measurement agreement. RESULTS: All but one parameter showed an excellent agreement (>80%) between both devices in the ICC analysis. No criteria for Bland-Altman LoA and bias were found in the literature regarding ECG parameters. CONCLUSION: The results of the ICC and Bland-Altman methods demonstrate that the BITalino system can be considered as an equivalent recording device for stationary ECG recordings in psychophysiological experiments.

A three-armed cognitive-motor exercise intervention to increase spatial orientation and life-space mobility in nursing home residents: study protocol of a randomized controlled trial in the PROfit project.

BACKGROUND: In nursing home residents, the combination of decreasing mobility and declining cognitive abilities, including spatial orientation, often leads to reduced physical activity (PA) and life-space (LS) mobility. As a consequence of sedentary behavior, there is a lack of social interaction and cognitive stimulation, resulting in low quality of life. It has not yet been examined whether cognitive-motor training including spatial cognitive tasks is suitable to improve spatial orientation and, as a consequence, to enlarge LS mobility, and increase well-being and general cognitive-motor functioning. Therefore, the overall goal of this multicentric randomized controlled trial (RCT) is to compare the effect of three different intervention approaches including functional exercise and orientation tasks on PA, LS and spatial orientation in nursing home residents. METHODS: A three-arm single-blinded multicenter RCT with a wait-list control group will be conducted in a sample of 513 individuals (needed according to power analysis) in three different regions in Germany. In each nursing home, one of three different intervention approaches will be delivered to participating residents for 12 weeks, twice a week for 45 min each: The PROfit basic group will perform functional strength, balance, flexibility, and walking exercises always at the same location, whereas the PROfit plus group changes the location three times while performing similar/the same exercises as the PROfit basic group. The PROfit orientation group receives navigation tasks in addition to the relocation during the intervention. Physical and cognitive functioning as well as psychological measures will be assessed in all study groups at baseline. Participants will then be randomized into either the intervention group or the wait-list control group. After 12 weeks, and after 24 weeks the measures will be repeated. DISCUSSION: This study evaluates whether the three different interventions are feasible to reduce the decline of or even improve PA, LS, and spatial orientation in nursing home residents. By adding different training locations in PROfit plus, the program is expected to be superior to PROfit basic in increasing physical and cognitive parameters. Moreover, we expect the PROfit orientation intervention to be most effective in terms of PA, LS, and spatial orientation due to two mechanisms: (1) increased physical and cognitive activity will enhance cognitive-motor capacity and (2) the spatial training will help to build up cognitive strategies to compensate for age-related loss of spatial orientation abilities and related limitations. TRIAL REGISTRATION: The trial was prospectively registered at DRKS.de with registration number DRKS00021423 on April 16, 2020 and was granted permission by the Technical University Berlin local ethics committee (No. GR_14_20191217).

Neuroadaptive technology enables implicit cursor control based on medial prefrontal cortex activity.

The effectiveness of today's human-machine interaction is limited by a communication bottleneck as operators are required to translate high-level concepts into a machine-mandated sequence of instructions. In contrast, we demonstrate effective, goal-oriented control of a computer system without any form of explicit communication from the human operator. Instead, the system generated the necessary input itself, based on real-time analysis of brain activity. Specific brain responses were evoked by violating the operators' expectations to varying degrees. The evoked brain activity demonstrated detectable differences reflecting congruency with or deviations from the operators' expectations. Real-time analysis of this activity was used to build a user model of those expectations, thus representing the optimal (expected) state as perceived by the operator. Based on this model, which was continuously updated, the computer automatically adapted itself to the expectations of its operator. Further analyses showed this evoked activity to originate from the medial prefrontal cortex and to exhibit a linear correspondence to the degree of expectation violation. These findings extend our understanding of human predictive coding and provide evidence that the information used to generate the user model is task-specific and reflects goal congruency. This paper demonstrates a form of interaction without any explicit input by the operator, enabling computer systems to become neuroadaptive, that is, to automatically adapt to specific aspects of their operator's mindset. Neuroadaptive technology significantly widens the communication bottleneck and has the potential to fundamentally change the way we interact with technology.

Two Independent Frontal Midline Theta Oscillations during Conflict Detection and Adaptation in a Simon-Type Manual Reaching Task.

One of the most firmly established factors determining the speed of human behavioral responses toward action-critical stimuli is the spatial correspondence between the stimulus and response locations. If both locations match, the time taken for response production is markedly reduced relative to when they mismatch, a phenomenon called the Simon effect. While there is a consensus that this stimulus-response (S-R) conflict is associated with brief (4-7 Hz) frontal midline theta (fmθ) complexes generated in medial frontal cortex, it remains controversial (1) whether there are multiple, simultaneously active theta generator areas in the medial frontal cortex that commonly give rise to conflict-related fmθ complexes; and if so, (2) whether they are all related to the resolution of conflicting task information. Here, we combined mental chronometry with high-density electroencephalographic measures during a Simon-type manual reaching task and used independent component analysis and time-frequency domain statistics on source-level activities to model fmθ sources. During target processing, our results revealed two independent fmθ generators simultaneously active in or near anterior cingulate cortex, only one of them reflecting the correspondence between current and previous S-R locations. However, this fmθ response is not exclusively linked to conflict but also to other, conflict-independent processes associated with response slowing. These results paint a detailed picture regarding the oscillatory correlates of conflict processing in Simon tasks, and challenge the prevalent notion that fmθ complexes induced by conflicting task information represent a unitary phenomenon related to cognitive control, which governs conflict processing across various types of response-override tasks.SIGNIFICANCE STATEMENT Humans constantly monitor their environment for and adjust their cognitive control settings in response to conflicts, an ability that arguably paves the way for survival in ever-changing situations. Anterior cingulate-generated frontal midline theta (fmθ) complexes have been hypothesized to play a role in this conflict-monitoring function. However, it remains a point of contention whether fmθ complexes govern conflict processing in a unitary, paradigm-nonspecific manner. Here, we identified two independent fmθ oscillations triggered during a Simon-type task, only one of them reflecting current and previous conflicts. Importantly, this signal differed in various respects (cortical origin, intertrial history) from fmθ phenomena in other response-override tasks, challenging the prevalent notion of conflict-induced fmθ as a unitary phenomenon associated with the resolution of conflict.

Prediction of cognitive conflict during unexpected robot behavior under different mental workload conditions in a physical human-robot collaboration.

Objective. Brain-computer interface (BCI) technology is poised to play a prominent role in modern work environments, especially a collaborative environment where humans and machines work in close proximity, often with physical contact. In a physical human robot collaboration (pHRC), the robot performs complex motion sequences. Any unexpected robot behavior or faulty interaction might raise safety concerns. Error-related potentials, naturally generated by the brain when a human partner perceives an error, have been extensively employed in BCI as implicit human feedback to adapt robot behavior to facilitate a safe and intuitive interaction. However, the integration of BCI technology with error-related potential for robot control demands failure-free integration of highly uncertain electroencephalography (EEG) signals, particularly influenced by the physical and cognitive state of the user. As a higher workload on the user compromises their access to cognitive resources needed for error awareness, it is crucial to study how mental workload variations impact the error awareness as it might raise safety concerns in pHRC. In this study, we aim to study how cognitive workload affects the error awareness of a human user engaged in a pHRC.Approach. We designed a blasting task with an abrasive industrial robot and manipulated the mental workload with a secondary arithmetic task of varying difficulty. EEG data, perceived workload, task and physical performance were recorded from 24 participants moving the robot arm. The error condition was achieved by the unexpected stopping of the robot in 33% of trials.Main results. We observed a diminished amplitude for the prediction error negativity (PEN) and error positivity (Pe), indicating reduced error awareness with increasing mental workload. We further observed an increased frontal theta power and increasing trend in the central alpha and central beta power after the unexpected robot stopping compared to when the robot stopped correctly at the target. We also demonstrate that a popular convolution neural network model, EEGNet, could predict the amplitudes of PEN and Pe from the EEG data prior to the error.Significance. This prediction model could be instrumental in developing an online prediction model that could forewarn the system and operators of the diminished error awareness of the user, alluding to a potential safety breach in error-related potential-based BCI system for pHRC. Therefore, our work paves the way for embracing BCI technology in pHRC to optimally adapt the robot behavior for personalized user experience using real-time brain activity, enriching the quality of the interaction.

Modeling the Intent to Interact with VR using Physiological Features.

OBJECTIVE: Mixed-Reality (XR) technologies promise a user experience (UX) that rivals the interactive experience with the real-world. The key facilitators in the design of such a natural UX are that the interaction has zero lag and that users experience no excess mental load. This is difficult to achieve due to technical constraints such as motion-to-photon latency as well as false-positives during gesture-based interaction.

Virtual Reality for Spatial Navigation.

Immersive virtual reality (VR) allows its users to experience physical space in a non-physical world. It has developed into a powerful research tool to investigate the neural basis of human spatial navigation as an embodied experience. The task of wayfinding can be carried out by using a wide range of strategies, leading to the recruitment of various sensory modalities and brain areas in real-life scenarios. While traditional desktop-based VR setups primarily focus on vision-based navigation, immersive VR setups, especially mobile variants, can efficiently account for motor processes that constitute locomotion in the physical world, such as head-turning and walking. When used in combination with mobile neuroimaging methods, immersive VR affords a natural mode of locomotion and high immersion in experimental settings, designing an embodied spatial experience. This in turn facilitates ecologically valid investigation of the neural underpinnings of spatial navigation.

Does Standing Up Enhance Performance on the Stroop Task in Healthy Young Adults? A Systematic Review and Meta-Analysis.

Understanding the changes in cognitive processing that accompany changes in posture can expand our understanding of embodied cognition and open new avenues for applications in (neuro)ergonomics. Recent studies have challenged the question of whether standing up alters cognitive performance. An electronic database search for randomized controlled trials was performed using Academic Search Complete, CINAHL Ultimate, MEDLINE, PubMed, and Web of Science following PRISMA guidelines, PICOS framework, and standard quality assessment criteria (SQAC). We pooled data from a total of 603 healthy young adults for incongruent and 578 for congruent stimuli and Stroop effect (mean age = 24 years). Using random-effects results, no difference was found between sitting and standing for the Stroop effect (Hedges' g = 0.13, 95% CI = -0.04 to 0.29, p = 0.134), even when comparing congruent (Hedges' g = 0.10; 95% CI: -0.132 to 0.339; Z = 0.86; p = 0.389) and incongruent (Hedges' g = 0.18; 95% CI: -0.072 to 0.422; Z = 1.39; p = 0.164) stimuli separately. Importantly, these results imply that changing from a seated to a standing posture in healthy young adults is unlikely to have detrimental effects on selective attention and cognitive control. To gain a full understanding of this phenomenon, further research should examine this effect in a population of healthy older adults, as well as in a population with pathology.

Using spontaneous eye blink-related brain activity to investigate cognitive load during mobile map-assisted navigation.

The continuous assessment of pedestrians' cognitive load during a naturalistic mobile map-assisted navigation task is challenging because of limited experimental control over stimulus presentation, human-map-interactions, and other participant responses. To overcome this challenge, the present study takes advantage of navigators' spontaneous eye blinks during navigation to serve as event markers in continuously recorded electroencephalography (EEG) data to assess cognitive load in a mobile map-assisted navigation task. We examined if and how displaying different numbers of landmarks (3 vs. 5 vs. 7) on mobile maps along a given route would influence navigators' cognitive load during navigation in virtual urban environments. Cognitive load was assessed by the peak amplitudes of the blink-related fronto-central N2 and parieto-occipital P3. Our results show increased parieto-occipital P3 amplitude indicating higher cognitive load in the 7-landmark condition, compared to showing 3 or 5 landmarks. Our prior research already demonstrated that participants acquire more spatial knowledge in the 5- and 7-landmark conditions compared to the 3-landmark condition. Together with the current study, we find that showing 5 landmarks, compared to 3 or 7 landmarks, improved spatial learning without overtaxing cognitive load during navigation in different urban environments. Our findings also indicate a possible cognitive load spillover effect during map-assisted wayfinding whereby cognitive load during map viewing might have affected cognitive load during goal-directed locomotion in the environment or vice versa. Our research demonstrates that users' cognitive load and spatial learning should be considered together when designing the display of future navigation aids and that navigators' eye blinks can serve as useful event makers to parse continuous human brain dynamics reflecting cognitive load in naturalistic settings.