Brain Networks Responsible for Sense of Agency: An EEG Study

BackgroundSelf-agency (SA) is a person’s feeling that his action was generated by himself. The neural substrates of SA have been investigated in many neuroimaging studies, but the functional connectivity of identified regions has rarely been investigated. The goal of this study is to investigate the neural network related to SA.MethodsSA of hand movements was modulated with virtual reality. We examined the cortical network relating to SA modulation with electroencephalography (EEG) power spectrum and phase coherence of alpha, beta, and gamma frequency bands in 16 right-handed, healthy volunteers.ResultsIn the alpha band, significant relative power changes and phase coherence of alpha band were associated with SA modulation. The relative power decrease over the central, bilateral parietal, and right temporal regions (C4, Pz, P3, P4, T6) became larger as participants more effectively controlled the virtual hand movements. The phase coherence of the alpha band within frontal areas (F7-FP2, F7-Fz) was directly related to changes in SA. The functional connectivity was lower as the participants felt that they could control their virtual hand. In the other frequency bands, significant phase coherences were observed in the frontal (or central) to parietal, temporal, and occipital regions during SA modulation (Fz-O1, F3-O1, Cz-O1, C3-T4L in beta band; FP1-T6, FP1-O2, F7-T4L, F8-Cz in gamma band).ConclusionsOur study suggests that alpha band activity may be the main neural oscillation of SA, which suggests that the neural network within the anterior frontal area may be important in the generation of SA.     

Optical Methods of Transformation of Visual Feedback

It should be noted that existing notions concerning the functioning of the oculomotor system (OMS) in human beings depend to a considerable degree on the methods of investigation chosen. Thus, the use of a number of new experimental procedures, for example, the presentation of "pulse" and double "pulse" stimuli [14, 15] and of stimuli stabilized relative to the retina [2, 5, 6] has yielded new data on the functioning of the OMS that by no means always agree with generally accepted notions.     

Differential Effects of Visual Feedback on Subjective Visual Vertical Accuracy and Precision

The brain constructs an internal estimate of the gravitational vertical by integrating multiple sensory signals. In darkness, systematic head-roll dependent errors in verticality estimates, as measured by the subjective visual vertical (SVV), occur. We hypothesized that visual feedback after each trial results in increased accuracy, as physiological adjustment errors (A−/E-effect) are likely based on central computational mechanisms and investigated whether such improvements were related to adaptational shifts of perceived vertical or to a higher cognitive strategy. We asked 12 healthy human subjects to adjust a luminous arrow to vertical in various head-roll positions (0 to 120deg right-ear down, 15deg steps). After each adjustment visual feedback was provided (lights on, display of previous adjustment and of an earth-vertical cross). Control trials consisted of SVV adjustments without feedback. At head-roll angles with the largest A-effect (90, 105, and 120deg), errors were reduced significantly (p<0.001) by visual feedback, i.e. roll under-compensation decreased, while precision of SVV was not significantly (p>0.05) influenced. In seven subjects an additional session with two consecutive blocks (first with, then without visual feedback) was completed at 90, 105 and 120deg head-roll. In these positions the error-reduction by the previous visual feedback block remained significant over the consecutive 18–24 min (post-feedback block), i.e., was still significantly (p<0.002) different from the control trials. Eleven out of 12 subjects reported having consciously added a bias to their perceived vertical based on visual feedback in order to minimize errors. We conclude that improvements of SVV accuracy by visual feedback, which remained effective after removal of feedback for ≥18 min, rather resulted from a cognitive strategy than by adapting the internal estimate of the gravitational vertical. The mechanisms behind the SVV therefore, remained stable, which is also supported by the fact that SVV precision – depending mostly on otolith input - was not affected by visual feedback.     

Re-Examining the Agentic Shift: The Sense of Agency Influences the Effectiveness of (Self)Persuasion

In the present study we investigated whether differences in the sense of agency influenced the effectiveness of both direct persuasion and self-persuasion techniques. By manipulating both the delay and contingency of the outcomes of actions, participants were led to experience either a low or high sense of agency. Participants were subsequently presented with arguments as to why a clean local environment is important (direct persuasion), or were asked to generate those arguments themselves (self-persuasion). Subsequently, participants’ cleanliness attitudes and willingness to participate in a campus cleanup were measured. The results show that techniques of direct persuasion influenced attitudes and volunteering behavior under conditions of low rather than high agency, whereas techniques of self-persuasion were most effective under conditions of high rather than low agency. The present findings therefore show how recent experiences of agency, a state based experience of control, can influence the effectiveness of both external and internal persuasion techniques.     

The Sense of Agency during Continuous Action: Performance Is More Important than Action-Feedback Association

The sense of agency refers to the feeling that one is controlling events through one’s own behavior. This study examined how task performance and the delay of events influence one’s sense of agency during continuous action accompanied by a goal. The participants were instructed to direct a moving dot into a square as quickly as possible by pressing the left and right keys on a keyboard to control the direction in which the dot traveled. The interval between the key press and response of the dot (i.e., direction change) was manipulated to vary task difficulty. Moreover, in the assisted condition, the computer ignored participants’ erroneous commands, resulting in improved task performance but a weaker association between the participants’ commands and actual movements of the dot relative to the condition in which all of the participants’ commands were executed (i.e., self-control condition). The results showed that participants’ sense of agency increased with better performance in the assisted condition relative to the self-control condition, even though a large proportion of their commands were not executed. We concluded that, when the action-feedback association was uncertain, cognitive inference was more dominant relative to the process of comparing predicted and perceived information in the judgment of agency.     

A Feedback Model of Attention and Context Dependence in Visual Cortical Networks

We have modeled biologically realistic neural networks that may be involved in contextual modulation of stimulus responses, as reported in the neurophysiological experiments of Motter (1994a, 1994b) (Journal of Neuroscience, 14:2179–2189 and 2190–2199). The networks of our model are structured hierarchically with feedforward, feedback, and lateral connections, totaling several thousand cells and about 300,000 synapses. The contextual modulation, arising from attention cues, is explicitly modeled as a feedback signal coming from the highest-order cortical network. The feedback signal arises from mutually inhibitory neurons with different stimulus preferences. Although our model is probably the simplest one consistent with available anatomical and physiological evidence and ignores the complexities that may exist in high-level cortical networks such as the prefrontal cortex, it reproduces the experimental results quite well and offers some guidance for future experiments. We also report the unexpected observation of 40 Hz oscillations in the model.     

The Efficiency of the Brain-Computer Interfaces Based on Motor Imagery with Tactile and Visual Feedback

In this study we compared tactile and visual feedbacks for the motor imagery-based brain–computer interface (BCI) in five healthy subjects. A vertical green bar from the center of the fixing cross to the edge of the screen was used as visual feedback. Vibration motors that were placed on the forearms of the right and the left hands and on the back of the subject’s neck were used as tactile feedback. A vibration signal was used to confirm the correct classification of the EEG patterns of the motor imagery of right and left hand movements and the rest task. The accuracy of recognition in the classification of the three states (right hand movement, left hand movement, and rest) in the BCI without feedback exceeded the random level (33% for the three states) for all the subjects and was rather high (67.8% ± 13.4% (mean ± standard deviation)). Including the visual and tactile feedback in the BCI did not significantly change the mean accuracy of recognition of mental states for all the subjects (70.5% ± 14.8% for the visual feedback and 65.9% ± 12.4% for the tactile feedback). The analysis of the dynamics of the movement imagery skill in BCI users with the tactile and visual feedback showed no significant differences between these types of feedback. Thus, it has been found that the tactile feedback can be used in the motor imagery-based BCI instead of the commonly used visual feedback, which greatly expands the possibilities of the practical application of the BCI.     

Modulation of Force below 1 Hz: Age-Associated Differences and the Effect of Magnified Visual Feedback

Oscillations in force output change in specific frequency bins and have important implications for understanding aging and pathological motor control. Although previous studies have demonstrated that oscillations from 0–1 Hz can be influenced by aging and visuomotor processing, these studies have averaged power within this bandwidth and not examined power in specific frequencies below 1 Hz. The purpose was to determine whether a differential modulation of force below 1 Hz contributes to changes in force control related to manipulation of visual feedback and aging. Ten young adults (25±4 yrs, 5 men) and ten older adults (71±5 yrs, 4 men) were instructed to accurately match a target force at 2% of their maximal isometric force for 35 s with abduction of the index finger. Visual feedback was manipulated by changing the visual angle (0.05°, 0.5°, 1.5°) or removing it after 15 s. Modulation of force below 1 Hz was quantified by examining the absolute and normalized power in seven frequency bins. Removal of visual feedback increased normalized power from 0–0.33 Hz and decreased normalized power from 0.66–1.0 Hz. In contrast, magnification of visual feedback (visual angles of 0.5° and 1.5°) decreased normalized power from 0–0.16 Hz and increased normalized power from 0.66–1.0 Hz. Older adults demonstrated a greater increase in the variability of force with magnification of visual feedback compared with young adults (P = 0.05). Furthermore, older adults exhibited differential force modulation of frequencies below 1 Hz compared with young adults (P<0.05). Specifically, older adults exhibited greater normalized power from 0–0.16 Hz and lesser normalized power from 0.66–0.83 Hz. The changes in force modulation predicted the changes in the variability of force with magnification of visual feedback (R² = 0.80). Our findings indicate that force oscillations below 1 Hz are associated with force control and are modified by aging and visual feedback.     

A Supplementary System for a Brain-Machine Interface Based on Jaw Artifacts for the Bidimensional Control of a Robotic Arm

Non-invasive Brain-Machine Interfaces (BMIs) are being used more and more these days to design systems focused on helping people with motor disabilities. Spontaneous BMIs translate user''s brain signals into commands to control devices. On these systems, by and large, 2 different mental tasks can be detected with enough accuracy. However, a large training time is required and the system needs to be adjusted on each session. This paper presents a supplementary system that employs BMI sensors, allowing the use of 2 systems (the BMI system and the supplementary system) with the same data acquisition device. This supplementary system is designed to control a robotic arm in two dimensions using electromyographical (EMG) signals extracted from the electroencephalographical (EEG) recordings. These signals are voluntarily produced by users clenching their jaws. EEG signals (with EMG contributions) were registered and analyzed to obtain the electrodes and the range of frequencies which provide the best classification results for 5 different clenching tasks. A training stage, based on the 2-dimensional control of a cursor, was designed and used by the volunteers to get used to this control. Afterwards, the control was extrapolated to a robotic arm in a 2-dimensional workspace. Although the training performed by volunteers requires 70 minutes, the final results suggest that in a shorter period of time (45 min), users should be able to control the robotic arm in 2 dimensions with their jaws. The designed system is compared with a similar 2-dimensional system based on spontaneous BMIs, and our system shows faster and more accurate performance. This is due to the nature of the control signals. Brain potentials are much more difficult to control than the electromyographical signals produced by jaw clenches. Additionally, the presented system also shows an improvement in the results compared with an electrooculographic system in a similar environment.     

Electroosmosis Dominates Electrophoresis of Antibiotic Transport Across the Outer Membrane Porin F

We report that the dynamics of antibiotic capture and transport across a voltage-biased OmpF nanopore is dominated by the electroosmotic flow rather than the electrophoretic force. By reconstituting an OmpF porin in an artificial lipid bilayer and applying an electric field across it, we are able to elucidate the permeation of molecules and their mechanism of transport. This field gives rise to an electrophoretic force acting directly on a charged substrate but also indirectly via coupling to all other mobile ions, causing an electroosmotic flow. The directionality and magnitude of this flow depends on the selectivity of the channel. Modifying the charge state of three different substrates (norfloxacin, ciprofloxacin, and enoxacin) by varying the pH between 6 and 9 while the charge and selectivity of OmpF is conserved allows us to work under conditions in which electroosmotic flow and electrophoretic forces add or oppose. This configuration allows us to identify and distinguish the contributions of the electroosmotic flow and the electrophoretic force on translocation. Statistical analysis of the resolvable dwell times reveals rich kinetic details regarding the direction and the stochastic movement of antibiotics inside the nanopore. We quantitatively describe the electroosmotic velocity component experienced by the substrates and their diffusion coefficients inside the porin with an estimate of the energy barrier experienced by the molecules caused by the interaction with the channel wall, which slows down the permeation by several orders of magnitude.     

Inter-Individual Difference in the Effect of Mirror Reflection-Induced Visual Feedback on Phantom Limb Awareness in Forearm Amputees

Objective

To test whether the phantom limb awareness could be altered by observing mirror reflection-induced visual feedback (MVF) in unilateral forearm amputees.

Methods

Ten unilateral forearm amputees were asked to perform bilateral (intact and phantom) synchronous wrist motions with and without MVF. During wrist motion, electromyographic activities in the extensor digitorum longus (EDL) and flexor carpi radialis muscles (FCR) were recorded with bipolar electrodes. Degree of wrist range of motion (ROM) was also recorded by electrogoniometry attached to the wrist joint of intact side. Subjects were asked to answer the degree of attainment of phantom limb motion using a visual analog scale (VAS: ranging from 0 (hard) to 10 (easy)).

Results

VAS and ROM were significantly increased by utilizing MVF, and the extent of an enhancement of the VAS and wrist ROM was positively correlated (r = 0.72, p<0.05). Although FCR EMG activity also showed significant enhancement by MVF, this was not correlated with the changes of VAS and ROM. Interestingly, while we found negative correlation between EDL EMG activity and wrist ROM, MVF generally affected to be increasing both EDL EMG and ROM.

Conclusions

Although there was larger extent of variability in the effect of MVF on phantom limb awareness, MVF has a potential to enhance phantom limb awareness, in case those who has a difficulty for the phantom limb motion. The present result suggests that the motor command to the missing limb can be re-activated by an appropriate therapeutic strategy such as mirror therapy.     

Automatic Imitation in Rhythmical Actions: Kinematic Fidelity and the Effects of Compatibility,Delay, and Visual Monitoring

We demonstrate that observation of everyday rhythmical actions biases subsequent motor execution of the same and of different actions, using a paradigm where the observed actions were irrelevant for action execution. The cycle time of the distractor actions was subtly manipulated across trials, and the cycle time of motor responses served as the main dependent measure. Although distractor frequencies reliably biased response cycle times, this imitation bias was only a small fraction of the modulations in distractor speed, as well as of the modulations produced when participants intentionally imitated the observed rhythms. Importantly, this bias was not only present for compatible actions, but was also found, though numerically reduced, when distractor and executed actions were different (e.g., tooth brushing vs. window wiping), or when the dominant plane of movement was different (horizontal vs. vertical). In addition, these effects were equally pronounced for execution at 0, 4, and 8 s after action observation, a finding that contrasts with the more short-lived effects reported in earlier studies. The imitation bias was also unaffected when vision of the hand was occluded during execution, indicating that this effect most likely resulted from visuomotor interactions during distractor observation, rather than from visual monitoring and guidance during execution. Finally, when the distractor was incompatible in both dimensions (action type and plane) the imitation bias was not reduced further, in an additive way, relative to the single-incompatible conditions. This points to a mechanism whereby the observed action’s impact on motor processing is generally reduced whenever this is not useful for motor planning. We interpret these findings in the framework of biased competition, where intended and distractor actions can be represented as competing and quasi-encapsulated sensorimotor streams.     

Microfibril Orientation Dominates the Microelastic Properties of Human Bone Tissue at the Lamellar Length Scale

The elastic properties of bone tissue determine the biomechanical behavior of bone at the organ level. It is now widely accepted that the nanoscale structure of bone plays an important role to determine the elastic properties at the tissue level. Hence, in addition to the mineral density, the structure and organization of the mineral nanoparticles and of the collagen microfibrils appear as potential key factors governing the elasticity. Many studies exist on the role of the organization of collagen microfibril and mineral nanocrystals in strongly remodeled bone. However, there is no direct experimental proof to support the theoretical calculations. Here, we provide such evidence through a novel approach combining several high resolution imaging techniques: scanning acoustic microscopy, quantitative scanning small-Angle X-ray scattering imaging and synchrotron radiation computed microtomography. We find that the periodic modulations of elasticity across osteonal bone are essentially determined by the orientation of the mineral nanoparticles and to a lesser extent only by the particle size and density. Based on the strong correlation between the orientation of the mineral nanoparticles and the collagen molecules, we conclude that the microfibril orientation is the main determinant of the observed undulations of microelastic properties in regions of constant mineralization in osteonal lamellar bone. This multimodal approach could be applied to a much broader range of fibrous biological materials for the purpose of biomimetic technologies.