Neural Coupling Research Articles

Development of an automatic rotational orthosis for walking with arm swing.

Interlimb neural coupling is often observed during normal gait and is postulated to be important for gait restoration. In order to provide a testbed for investigation of interlimb neural coupling, we previously developed a rotational orthosis for walking with arm swing (ROWAS). The present study aimed to develop and evaluate the feasibility of a new system, viz. an automatic ROWAS (aROWAS). We developed the mechanical structures of aROWAS in SolidWorks, and implemented the concept in a prototype. Normal gait data from walking at various speeds were used as reference trajectories of the shoulder, hip, knee and ankle joints. The aROWAS prototype was tested in three able-bodied subjects. The prototype could automatically adjust to size and height, and automatically produced adaptable coordinated performance in the upper and lower limbs, with joint profiles similar to those occurring in normal gait. The subjects reported better acceptance in aROWAS than in ROWAS. The aROWAS system was deemed feasible among able-bodied subjects.

Neural Cross-Frequency Coupling Functions

Although neural interactions are usually characterized only by their coupling strength and directionality, there is often a need to go beyond this by establishing the functional mechanisms of the interaction. We introduce the use of dynamical Bayesian inference for estimation of the coupling functions of neural oscillations in the presence of noise. By grouping the partial functional contributions, the coupling is decomposed into its functional components and its most important characteristics—strength and form—are quantified. The method is applied to characterize the δ-to-α phase-to-phase neural coupling functions from electroencephalographic (EEG) data of the human resting state, and the differences that arise when the eyes are either open (EO) or closed (EC) are evaluated. The δ-to-α phase-to-phase coupling functions were reconstructed, quantified, compared, and followed as they evolved in time. Using phase-shuffled surrogates to test for significance, we show how the strength of the direct coupling, and the similarity and variability of the coupling functions, characterize the EO and EC states for different regions of the brain. We confirm an earlier observation that the direct coupling is stronger during EC, and we show for the first time that the coupling function is significantly less variable. Given the current understanding of the effects of e.g., aging and dementia on δ-waves, as well as the effect of cognitive and emotional tasks on α-waves, one may expect that new insights into the neural mechanisms underlying certain diseases will be obtained from studies of coupling functions. In principle, any pair of coupled oscillations could be studied in the same way as those shown here.

3D printing scaffold coupled with low level light therapy for neural tissue regeneration

3D printing has shown promise for neural regeneration by providing customized nerve scaffolds to structurally support and bridge the defect gap as well as deliver cells or various bioactive substances. Low-level light therapy (LLLT) exhibits positive effects on rehabiliation of degenerative nerves and neural disorders. With this in mind, we postulate that 3D printed neural scaffold coupling with LLLT will generate a new strategy to repair neural degeneration. To achieve this goal, we applied red laser light to stimualte neural stem cells on 3D printed scaffolds and investigated the subsequent cell response with respect to cell proliferation and differentiation. Here we show that cell prolifeartion rate and intracellular reactive oxgen species synthesis were significantly increased after 15 s laser stimulation follwed by 1 d culture. Over culturing time of 14 d in vitro, the laser stimulation promoted neuronal differentiation of neural stem cells, while the glial differentiation was suppressed based on results of both immunocytochemistry studies and real-time quantitative reverse transcription polymerase chain reaction testing. These findings suggest that integration of 3D printing and LLLT might provide a powerful methodology for neural tissue engineering.

Perception of social synchrony induces mother-child gamma coupling in the social brain.

The recent call to move from focus on one brain’s functioning to two-brain communication initiated a search for mechanisms that enable two humans to coordinate brain response during social interactions. Here, we utilized the mother–child context as a developmentally salient setting to study two-brain coupling. Mothers and their 9-year-old children were videotaped at home in positive and conflictual interactions. Positive interactions were microcoded for social synchrony and conflicts for overall dialogical style. Following, mother and child underwent magnetoencephalography while observing the positive vignettes. Episodes of behavioral synchrony, compared to non-synchrony, increased gamma-band power in the superior temporal sulcus (STS), hub of social cognition, mirroring and mentalizing. This neural pattern was coupled between mother and child. Brain-to-brain coordination was anchored in behavioral synchrony; only during episodes of behavioral synchrony, but not during non-synchronous moments, mother’s and child's STS gamma power was coupled. Importantly, neural synchrony was not found during observation of unfamiliar mother-child interaction Maternal empathic/dialogical conflict style predicted mothers’ STS activations whereas child withdrawal predicted attenuated STS response in both partners. Results define a novel neural marker for brain-to-brain synchrony, highlight the role of rapid bottom-up oscillatory mechanisms for neural coupling and indicate that behavior-based processes may drive synchrony between two brains during social interactions.

Concurrent tACS-fMRI Reveals Causal Influence of Power Synchronized Neural Activity on Resting State fMRI Connectivity.

Resting state fMRI (rs-fMRI) is commonly used to study the brain's intrinsic neural coupling, which reveals specific spatiotemporal patterns in the form of resting state networks (RSNs). It has been hypothesized that slow rs-fMRI oscillations (<0.1 Hz) are driven by underlying electrophysiological rhythms that typically occur at much faster timescales (>5 Hz); however, causal evidence for this relationship is currently lacking. Here we measured rs-fMRI in humans while applying transcranial alternating current stimulation (tACS) to entrain brain rhythms in left and right sensorimotor cortices. The two driving tACS signals were tailored to the individual's α rhythm (8-12 Hz) and fluctuated in amplitude according to a 1 Hz power envelope. We entrained the left versus right hemisphere in accordance to two different coupling modes where either α oscillations were synchronized between hemispheres (phase-synchronized tACS) or the slower oscillating power envelopes (power-synchronized tACS). Power-synchronized tACS significantly increased rs-fMRI connectivity within the stimulated RSN compared with phase-synchronized or no tACS. This effect outlasted the stimulation period and tended to be more effective in individuals who exhibited a naturally weak interhemispheric coupling. Using this novel approach, our data provide causal evidence that synchronized power fluctuations contribute to the formation of fMRI-based RSNs. Moreover, our findings demonstrate that the brain's intrinsic coupling at rest can be selectively modulated by choosing appropriate tACS signals, which could lead to new interventions for patients with altered rs-fMRI connectivity.SIGNIFICANCE STATEMENT Resting state fMRI (rs-fMRI) has become an important tool to estimate brain connectivity. However, relatively little is known about how slow hemodynamic oscillations measured with fMRI relate to electrophysiological processes. It was suggested that slowly fluctuating power envelopes of electrophysiological signals synchronize across brain areas and that the topography of this activity is spatially correlated to resting state networks derived from rs-fMRI. Here we take a novel approach to address this problem and establish a causal link between the power fluctuations of electrophysiological signals and rs-fMRI via a new neuromodulation paradigm, which exploits these power synchronization mechanisms. These novel mechanistic insights bridge different scientific domains and are of broad interest to researchers in the fields of Medical Imaging, Neuroscience, Physiology, and Psychology.

Abnormal frontoparietal synaptic gain mediating the P300 in patients with psychotic disorder and their unaffected relatives.

The "dysconnection hypothesis" of psychosis suggests that a disruption of functional integration underlies cognitive deficits and clinical symptoms. Impairments in the P300 potential are well documented in psychosis. Intrinsic (self-)connectivity in a frontoparietal cortical hierarchy during a P300 experiment was investigated. Dynamic Causal Modeling was used to estimate how evoked activity results from the dynamics of coupled neural populations and how neural coupling changes with the experimental factors. Twenty-four patients with psychotic disorder, twenty-four unaffected relatives, and twenty-five controls underwent EEG recordings during an auditory oddball paradigm. Sixteen frontoparietal network models (including primary auditory, superior parietal, and superior frontal sources) were analyzed and an optimal model of neural coupling, explaining diagnosis and genetic risk effects, as well as their interactions with task condition were identified. The winning model included changes in connectivity at all three hierarchical levels. Patients showed decreased self-inhibition-that is, increased cortical excitability-in left superior frontal gyrus across task conditions, compared with unaffected participants. Relatives had similar increases in excitability in left superior frontal and right superior parietal sources, and a reversal of the normal synaptic gain changes in response to targets relative to standard tones. It was confirmed that both subjects with psychotic disorder and their relatives show a context-independent loss of synaptic gain control at the highest hierarchy levels. The relatives also showed abnormal gain modulation responses to task-relevant stimuli. These may be caused by NMDA-receptor and/or GABAergic pathologies that change the excitability of superficial pyramidal cells and may be a potential biological marker for psychosis. Hum Brain Mapp 38:3262-3276, 2017. © 2017 Wiley Periodicals, Inc.

SU72. Abnormal Frontal Synaptic Gain Mediating the P300 in Patients With Psychosis and Their Unaffected Relatives

Abstract Background: The “dysconnection hypothesis” of psychosis suggests that a disruption of functional integration underlies cognitive deficits and clinical symptoms. Impairments in the P300 potential are well documented in psychosis. We investigated intrinsic (self-)connectivity in a cortical hierarchy during a P300 experiment. We used Dynamic Causal Modeling to estimate how evoked activity results from the dynamics of coupled neural populations and how neural coupling changes with the experimental factors. Methods: Twenty-four patients with psychosis, 24 unaffected relatives, and 25 controls underwent EEG recordings during an auditory oddball paradigm. We analyzed 16 frontoparietal models (primary auditory, superior parietal, and superior frontal sources) and identified an optimal model of neural coupling, explaining diagnosis and genetic risk effects, as well as their interactions with task condition. Results: The winning model included changes in connectivity at all 3 hierarchical levels. Patients showed decreased self-inhibition (ie, increased cortical excitability) in left superior frontal gyrus across task conditions, compared to unaffected participants. Relatives had similar increases in excitability in left superior frontal and right superior parietal sources, and a reversal of the normal synaptic gain changes in response to targets, relative to standard tones. Conclusion: We confirmed that both subjects with psychosis and their relatives show a context-independent loss of synaptic gain control at the highest levels of the hierarchy. The relatives also showed abnormal gain modulation responses to task-relevant stimuli. These may be caused by NMDA-receptor and/or GABAergic pathologies that change the excitability of superficial pyramidal cells and may be a potential biological marker for psychosis.

Measuring speaker\u2013listener neural coupling with functional near infrared spectroscopy

The present study investigates brain-to-brain coupling, defined as inter-subject correlations in the hemodynamic response, during natural verbal communication. We used functional near-infrared spectroscopy (fNIRS) to record brain activity of 3 speakers telling stories and 15 listeners comprehending audio recordings of these stories. Listeners’ brain activity was significantly correlated with speakers’ with a delay. This between-brain correlation disappeared when verbal communication failed. We further compared the fNIRS and functional Magnetic Resonance Imaging (fMRI) recordings of listeners comprehending the same story and found a significant relationship between the fNIRS oxygenated-hemoglobin concentration changes and the fMRI BOLD in brain areas associated with speech comprehension. This correlation between fNIRS and fMRI was only present when data from the same story were compared between the two modalities and vanished when data from different stories were compared; this cross-modality consistency further highlights the reliability of the spatiotemporal brain activation pattern as a measure of story comprehension. Our findings suggest that fNIRS can be used for investigating brain-to-brain coupling during verbal communication in natural settings.

Development and Feasibility Assessment of a Rotational Orthosis for Walking with Arm Swing.

Interlimb neural coupling might underlie human bipedal locomotion, which is reflected in the fact that people swing their arms synchronously with leg movement in normal gait. Therefore, arm swing should be included in gait training to provide coordinated interlimb performance. The present study aimed to develop a Rotational Orthosis for Walking with Arm Swing (ROWAS), and evaluate its feasibility from the perspectives of implementation, acceptability and responsiveness. We developed the mechanical structures of the ROWAS system in SolidWorks, and implemented the concept in a prototype. Normal gait data were used as the reference performance of the shoulder, hip, knee and ankle joints of the prototype. The ROWAS prototype was tested for function assessment and further evaluated using five able-bodied subjects for user feedback. The ROWAS prototype produced coordinated performance in the upper and lower limbs, with joint profiles similar to those occurring in normal gait. The subjects reported a stronger feeling of walking with arm swing than without. The ROWAS system was deemed feasible according to the formal assessment criteria.

Neuromodulation in response to electroconvulsive therapy in schizophrenia and major depression

BackgroundElectroconvulsive therapy (ECT) is one of the most effective treatments in severe and treatment-resistant major depressive disorder (MDD). ECT has been also shown to be effective in schizophrenia (SZ), particularly when rapid symptom reduction is needed or in cases of resistance to drug-treatment. However, its precise mechanisms of action remain largely unknown. Objective/HypothesisThis study examined whether ECT exerts disorder-specific or unspecific modulation of brain structure and function in SZ and MDD. MethodsWe investigated neuromodulatory effects of right-sided unilateral ECT in pharmacoresistant patients with SZ or MDD. Magnetic resonance imaging was conducted before and after ECT to investigate treatment-related effects on brain structure and function. Imaging data were analyzed by means of Voxel Based Morphometry and Resting State Functional Connectivity (RSFC) methods. ResultsRight unilateral ECT induced transdiagnostic regional increases of limbic gray matter and modulations of neural coupling at rest. Structural effects were accompanied by a decrease in RSFC within temporoparietal, prefrontal and cortical midline structures, and an increase in hypothalamic RSFC. The extent of structural and functional change was partially inversely associated with the baseline measures. ConclusionThe present findings provide first evidence for transdiagnostic changes of brain structure together with modulation of brain function after ECT. The data indicate diagnosis-unspecific mechanisms of action with respect to regional gray matter volume and resting-state functional connectivity.

Human Exploration of Enclosed Spaces through Echolocation.

Some blind humans have developed echolocation, as a method of navigation in space. Echolocation is a truly active sense because subjects analyze echoes of dedicated, self-generated sounds to assess space around them. Using a special virtual space technique, we assess how humans perceive enclosed spaces through echolocation, thereby revealing the interplay between sensory and vocal-motor neural activity while humans perform this task. Sighted subjects were trained to detect small changes in virtual-room size analyzing real-time generated echoes of their vocalizations. Individual differences in performance were related to the type and number of vocalizations produced. We then asked subjects to estimate virtual-room size with either active or passive sounds while measuring their brain activity with fMRI. Subjects were better at estimating room size when actively vocalizing. This was reflected in the hemodynamic activity of vocal-motor cortices, even after individual motor and sensory components were removed. Activity in these areas also varied with perceived room size, although the vocal-motor output was unchanged. In addition, thalamic and auditory-midbrain activity was correlated with perceived room size; a likely result of top-down auditory pathways for human echolocation, comparable with those described in echolocating bats. Our data provide evidence that human echolocation is supported by active sensing, both behaviorally and in terms of brain activity. The neural sensory-motor coupling complements the fundamental acoustic motor-sensory coupling via the environment in echolocation.SIGNIFICANCE STATEMENT Passive listening is the predominant method for examining brain activity during echolocation, the auditory analysis of self-generated sounds. We show that sighted humans perform better when they actively vocalize than during passive listening. Correspondingly, vocal motor and cerebellar activity is greater during active echolocation than vocalization alone. Motor and subcortical auditory brain activity covaries with the auditory percept, although motor output is unchanged. Our results reveal behaviorally relevant neural sensory-motor coupling during echolocation.

Interindividual differences in motor network connectivity and behavioral response to iTBS in stroke patients.

Cerebral plasticity-inducing approaches like repetitive transcranial magnetic stimulation (rTMS) are of high interest in situations where reorganization of neural networks can be observed, e.g., after stroke. However, an increasing number of studies suggest that improvements in motor performance of the stroke-affected hand following modulation of primary motor cortex (M1) excitability by rTMS shows a high interindividual variability. We here tested the hypothesis that in stroke patients the interindividual variability of behavioral response to excitatory rTMS is related to interindividual differences in network connectivity of the stimulated region. Chronic stroke patients (n = 14) and healthy controls (n = 12) were scanned with functional magnetic resonance imaging (fMRI) while performing a simple hand motor task. Dynamic causal modeling (DCM) was used to investigate effective connectivity of key motor regions. On two different days after the fMRI experiment, patients received either intermittent theta-burst stimulation (iTBS) over ipsilesional M1 or control stimulation over the parieto-occipital cortex. Motor performance and TMS parameters of cortical excitability were measured before and after iTBS. Our results revealed that patients with better motor performance of the affected hand showed stronger endogenous coupling between supplemental motor area (SMA) and M1 before starting the iTBS intervention. Applying iTBS to ipsilesional M1 significantly increased ipsilesional M1 excitability and decreased contralesional M1 excitability as compared to control stimulation. Individual behavioral improvements following iTBS specifically correlated with neural coupling strengths in the stimulated hemisphere prior to stimulation, especially for connections targeting the stimulated M1. Combining endogenous connectivity and behavioral parameters explained 82% of the variance in hand motor performance observed after iTBS. In conclusion, the data suggest that the individual susceptibility to iTBS after stroke is influenced by interindividual differences in motor network connectivity of the lesioned hemisphere.

Cortical interactions during the resolution of information processing demands in autism spectrum disorders

IntroductionOur flexible and adaptive interactions with the environment are guided by our individual representation of the physical world, estimated through sensation and evaluation of available information against prior knowledge. When linking sensory evidence with higher‐level expectations for action, the central nervous system (CNS) in typically developing (TD) individuals relies in part on distributed and interacting cortical regions to communicate neuronal signals flexibly across the brain. Increasing evidence suggests that the balance between levels of signal and noise during information processing may be disrupted in individuals with Autism Spectrum Disorders (ASD).MethodsParticipants with and without ASD performed a visuospatial interference task while undergoing functional Magnetic Resonance Imaging (fMRI). We empirically estimated parameters characterizing participants’ latencies and their subtle fluctuations (noise accumulation) over the 16‐min scan. We modeled hemodynamic activation and used seed‐based analyses of neural coupling to study dysfunction in interference‐specific connectivity in a subset of ASD participants who were nonparametrically matched to TD participants on age, male‐to‐female ratio, and magnitude of movement during the scan.ResultsStochastic patterns of response fluctuations reveal significantly higher noise‐to‐signal levels and a more random and noisy structure in ASD versus TD participants, and in particular ASD adults who have the greatest clinical autistic deficits. While individuals with ASD show an overall weaker modulation of interference‐specific functional connectivity relative to TD individuals, in particular between the seeds of Anterior Cingulate Cortex (ACC) and Inferior Parietal Sulcus (IPS) and the rest of the brain, we found that in ASD, higher uncertainty during the task is linked to increased interference‐specific coupling between bilateral anterior insula and prefrontal cortex.ConclusionsSubtle and informative differences in the structure of experiencing information exist between ASD and TD individuals. Our findings reveal in ASD an atypical capacity to apply previously perceived information in a manner optimal for adaptive functioning, plausibly revealing suboptimal message‐passing across the CNS.

Motor adaptations to unilateral quadriceps fatigue during a bilateral pedaling task.

This study was designed to investigate how motor coordination adapts to unilateral fatigue of the quadriceps during a constant-load bilateral pedaling task. We first hypothesized that this local fatigue would not be compensated within the fatigued muscles leading to a decreased knee extension power. Then, we aimed to determine whether this decrease would be compensated by between-joints compensations within the ipsilateral leg and/or an increased contribution of the contralateral leg. Fifteen healthy volunteers were tested during pedaling at 350W before and after a fatigue protocol consisting of 15minutes of electromyostimulation on the quadriceps muscle. Motor coordination was assessed from myoelectrical activity (22 muscles) and joint powers calculated through inverse dynamics. Maximal knee extension torque decreased by 28.3%±6.8% (P<.0005) immediately after electromyostimulation. A decreased knee extension power produced by the ipsilateral leg was observed during pedaling (-22.8±12.3W, -17.0%±9.4%; P<.0005). To maintain the task goal, participants primarily increased the power produced by the non-fatigued contralateral leg during the flexion phase. This was achieved by an increase in hip flexion power confirmed by a higher activation of the tensor fascia latae. These results suggest no adjustment of neural drive to the fatigued muscles and demonstrate no concurrent ipsilateral compensation by the non-fatigued muscles involved in the extension pedaling phase. Although interindividual variability was observed, findings provide evidence that participants predominantly adapted by compensating with the contralateral leg during its flexion phase. Both neural (between legs) and mechanical (between pedals) couplings and the minimization of cost functions might explain these results.

Rhythmic wrist movements facilitate the soleus H-reflex and non-voluntary air-stepping in humans

Neural coupling between the upper and lower limbs during human walking is supported by modulation of cross-limb reflexes and the presence of rhythmic activity in the proximal arm muscles. Nevertheless, the involvement of distal arm muscles in cyclic movements and sensorimotor neuromodulation is also suggested given their step-synchronized activation in many locomotor-related tasks (e.g., swimming, skiing, climbing, cycling, crawling, etc.). Here we investigated the effect of rhythmic wrist movements, separately and in conjunction with arm swinging, on the characteristics of non-voluntary cyclic leg movements evoked by muscle vibration in a gravity neutral position and on the soleus H-reflex of the stationary legs. For the H-reflex modulation, five conditions were compared: stationary arms, voluntary alternating upper limb swinging, combined upper limb and wrist motion, wrist movements only and motion of the upper limbs with addition of load. Rhythmic wrist movements significantly facilitated the amplitude of non-voluntary leg oscillations, including ankle joint oscillations, and the H-reflex. The latter effect was related to rhythmicity of wrist motion rather than to a simple extra tension in the upper limb muscles (a kind of the Jendrassik manoeuvre) since adding resistance to arm oscillations (without flexion-extension in the wrist joint) had an opposite inhibitory effect on the H-reflex. Our results further support the existence of connections between the distal parts of the upper and lower extremities at the neural level, suggesting that wrist joint movements can be an important component of motor neurorehabilitation.

Neural coupling between homologous muscles during bimanual tasks: effects of visual and somatosensory feedback.

This study investigated the effects of somatosensory feedback during bimanual tasks on the neural coupling between arm muscles, which remains largely unexplored. Somatosensory feedback using a balancing apparatus, compared with visual feedback, significantly increased neural coupling between homologous muscles (indicated by intermuscular coherence values) and improved temporal correlation of bilateral force production. Notably, feedback type modulated coherence in the α- and γ-bands (more subcortical pathways), whereas task type mainly affected β-band coherence (corticospinal pathway).

Neural coupling of cooperative hand movements after stroke: role of ipsilateral afference.

We investigated the role of ipsilateral ascending pathways in the neural coupling underlying cooperative hand movements of stroke subjects. Ipsi‐ and contralateral somatosensory evoked potentials (SSEP) were recorded following ulnar nerve stimulation during cooperative and non‐cooperative hand movements. The amplitude ratio, that is, ipsilateral divided by contralateral amplitude, was highest during the cooperative task when the affected arm was stimulated, reflecting an enhanced afferent volley to the unaffected hemisphere. The presence of ipsilateral SSEP from the paretic arm was closely related with the patients' hand function. This shows for the first time a laterality in ascending pathways after unilateral stroke and implies an involvement of the unaffected hemisphere in the control of paretic hand movements.

Weak electric fields detectability in a noisy neural network.

We investigate the detectability of weak electric field in a noisy neural network based on Izhikevich neuron model systematically. The neural network is composed of excitatory and inhibitory neurons with similar ratio as that in the mammalian neocortex, and the axonal conduction delays between neurons are also considered. It is found that the noise intensity can modulate the detectability of weak electric field. Stochastic resonance (SR) phenomenon induced by white noise is observed when the weak electric field is added to the network. It is interesting that SR almost disappeared when the connections between neurons are cancelled, suggesting the amplification effects of the neural coupling on the synchronization of neuronal spiking. Furthermore, the network parameters, such as the connection probability, the synaptic coupling strength, the scale of neuron population and the neuron heterogeneity, can also affect the detectability of the weak electric field. Finally, the model sensitivity is studied in detail, and results show that the neural network model has an optimal region for the detectability of weak electric field signal.

Adaptation of horizontal eye alignment in the presence of prism in young children.

Young children experience decreased convergence and increased accommodation demands relative to adults, as a result of their small interpupillary distance and hyperopic refraction. Those with typical amounts of hyperopic refractive error must accommodate more than an emmetrope to achieve focused retinal images, which may also drive additional convergence through the neural coupling. Adults and older children have demonstrated vergence adaptation to a variety of visual stimuli. Can vergence adaptation help younger children achieve alignment in the presence of these potentially conflicting demands? Purkinje image eye tracking and eccentric photorefraction were used to record simultaneous vergence and accommodation responses in adults and young children (3–6 years). To assess vergence adaptation, heterophoria was monitored before, during, and after adaptation induced by both base-in and base-out prisms. Adaptation was observed in both adults and young children with no significant effect of age, F(1, 34) = 0.014, p = 0.907. Changes in accommodation between before, during, and after adaptation were less than 0.5 D in binocular viewing. Typically developing children appear capable of vergence adaptation, which might play an important role in the maintenance of eye alignment under their changing visual demands.

Neural correlates of maintaining generated images in visual working memory.

How are images that have been assembled from their constituting elements maintained as a coherent representation in visual working memory (vWM)? Here, we compared two conditions of vWM maintenance that only differed in how vWM contents had been created. Participants maintained images that they either had to assemble from single features or that they had perceived as complete objects. Object complexity varied between two and four features. We analyzed electroencephalogram phase coupling as a measure of cortical connectivity in a time interval immediately before a probe stimulus appeared. We assumed that during this time both groups maintained essentially the same images, but that images constructed from their elements would require more neural coupling than images based on a complete percept. Increased coupling between frontal and parietal-to-occipital cortical sources was found for the maintenance of constructed in comparison to nonconstructed objects in the theta, alpha, beta, and gamma frequency bands. A similar pattern was found for an increase in vWM load (2 vs. 4 features) for nonconstructed objects. Under increased construction load (2 vs. 4 features for constructed images), the pattern was restricted to fronto-parietal couplings, suggesting that the fronto-parietal attention network is coping with the higher attentional demands involved in maintaining constructed images, but without increasing the communication with the occipital visual buffer in which the visual representations are assumed to be stored. We conclude from these findings that the maintenance of constructed images in vWM requires additional attentional processes to keep object elements together as a coherent representation. Hum Brain Mapp 37:4349-4362, 2016. © 2016 Wiley Periodicals, Inc.