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Neuronal signals encoding the animal's position widely modulate neocortical processing. While these signals are assumed to depend on hippocampal output, their origin has not been investigated directly. Here, we asked which brain region sends position information to the retrosplenial cortex (RSC), a key circuit for memory and navigation. We comprehensively characterized the long-range inputs to agranular RSC using two-photon axonal imaging in head-fixed mice performing a spatial task in darkness. Surprisingly, most long-range pathways convey position information, but with notable differences. Axons from the secondary motor and posterior parietal cortex transmit the most position information. By contrast, axons from the anterior cingulate and orbitofrontal cortex and thalamus convey substantially less position information. Axons from the primary and secondary visual cortex contribute negligibly. This demonstrates that the hippocampus is not the only source of position information. Instead, the RSC is a hub in a distributed brain network that shares position information.

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BACKGROUND: Neuroimaging and lesion studies suggested that the dorsolateral prefrontal and posterior parietal cortices mediate visual metacognitive awareness. The causal evidence provided by non-invasive brain stimulation, however, is inconsistent. OBJECTIVE/HYPOTHESIS: Here we revisit a major figure discrimination experiment adding a new Kanizsa figure task trying to resolve whether bilateral continuous theta-burst transcranial magnetic stimulation (cTBS) over these regions affects perceptual metacognition. Specifically, we tested whether subjective visibility ratings and/or metacognitive efficiency are lower when cTBS is applied to these two regions in comparison to an active control region. METHODS: A within-subjects design including three sessions spaced by one-week intervals was implemented. In each session, every participant was administered bilateral cTBS to either prefrontal, control or parietal cortices. Two concurrent tasks were performed, a real and an illusory figure task, stabilising objective performance with use of an adaptive staircase procedure. RESULTS: When performing the replicated task, cTBS was found insufficient to disrupt neither visibility ratings nor metacognitive efficiency. However, with use of Kanizsa style illusory figures, cTBS over the dorsolateral prefrontal, but not over the posterior parietal cortex, was observed to significantly diminish metacognitive efficiency. CONCLUSION(S): Real and illusory figure tasks demonstrated different cTBS effects. A possible explanation is the involvement of the prefrontal cortex in the creation of expectations, which is necessary for efficient metacognition. Failure to replicate previous findings for the real figure task, however, cannot be said to support, conclusively, the notion that these brain regions have a causal role in metacognitive awareness. This inconsistent finding may result from certain limitations of our study, thereby suggesting the need for yet further investigation.

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Objective. Enable neural control of individual prosthetic fingers for participants with upper-limb paralysis.Approach. Two tetraplegic participants were each implanted with a 96-channel array in the left posterior parietal cortex (PPC). One of the participants was additionally implanted with a 96-channel array near the hand knob of the left motor cortex (MC). Across tens of sessions, we recorded neural activity while the participants attempted to move individual fingers of the right hand. Offline, we classified attempted finger movements from neural firing rates using linear discriminant analysis with cross-validation. The participants then used the neural classifier online to control individual fingers of a brain-machine interface (BMI). Finally, we characterized the neural representational geometry during individual finger movements of both hands.Main Results. The two participants achieved 86% and 92% online accuracy during BMI control of the contralateral fingers (chance = 17%). Offline, a linear decoder achieved ten-finger decoding accuracies of 70% and 66% using respective PPC recordings and 75% using MC recordings (chance = 10%). In MC and in one PPC array, a factorized code linked corresponding finger movements of the contralateral and ipsilateral hands.Significance. This is the first study to decode both contralateral and ipsilateral finger movements from PPC. Online BMI control of contralateral fingers exceeded that of previous finger BMIs. PPC and MC signals can be used to control individual prosthetic fingers, which may contribute to a hand restoration strategy for people with tetraplegia.

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Attention includes three different functional components: generating and maintaining an alert state (alerting), orienting to sensory events (orienting), and resolving conflicts between alternative actions (executive control). Neuroimaging and patient studies suggest that the posterior parietal cortex (PPC) is involved in all three attention components. Transcranial magnetic stimulation (TMS) has repeatedly been applied over the PPC to study its functional role for shifts and maintenance of visuospatial attention. Most TMS-PPC studies used only detection tasks or orienting paradigms to investigate TMS-PPC effects on attention processes, neglecting the alerting and executive control components of attention. The objective of the present study was to investigate the role of PPC in all three functional components of attention: alerting, orienting, and executive control. To this end, we disrupted PPC with TMS (continuous theta-burst stimulation), to modulate subsequent performance on the Lateralized-Attention Network Test, used to assess the three attention components separately. Our results revealed hemifield-specific effects on alerting and executive control functions, but we did not find stimulation effects on orienting performance. While this field of research and associated clinical development have been predominantly focused on orienting performance, our results suggest that parietal cortex and its modulation may affect other aspects of attention as well.

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Background: Recovery of walking post-stroke is highly variable. Accurately measuring and documenting functional brain activation characteristics during walking can help guide rehabilitation. Previous work in this area has been limited to investigations of frontal brain regions and have not utilized recent technological and analytical advances for more accurate measurements. There were three aims for this study: to characterize the hemodynamic profile during walking post-stroke, to investigate regional changes in brain activation during different phases of walking, and to related brain changes to clinical measures. Methods: Functional near-infrared spectroscopy (fNIRS) along the pre-frontal, premotor, sensorimotor, and posterior parietal cortices was used on twenty individuals greater than six months post-stroke. Individual fNIRS optodes were digitized and used to estimate channel locations on each participant and short separation channels were used to control for extracerebral hemodynamic changes. Participants walked at their comfortable pace several times along a hallway while brain activation was recorded. Exploratory cluster analysis was conducted to determine if there was a link between brain activation and clinical measures. Results: Sustained activation was observed in the pre-frontal cortex with the ipsilesional hemisphere showing greater activation compared to the contralesional side. Sensorimotor cortex was active during the early, acceleration stage of walking only. Posterior parietal cortex showed changes in activation during the later, steady-state stage of walking. Faster gait speeds also related to increased activation in contralesional sensorimotor and posterior parietal cortices. Exploratory analysis clustered participants into two distinct groups based on their brain activation profiles and generally showed that individuals with greater activation tended to have better physical outcomes. Conclusions: These findings can guide future research for obtaining adequate power and determining factors that can be used as effect modifiers to reduce inter-subject variability. Overall, this is the first study to report specific oxygenated and deoxygenated hemoglobin changes in frontal to parietal regions during walking in the stroke population. Our results shed light on the importance of measuring brain activation across the cortex and show the importance of pre-frontal, sensorimotor, and posterior parietal cortices in walking after a stroke.

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Background: Walking within the community requires the ability to walk while simultaneously completing other tasks. After a stroke, completing an additional task while walking is significantly impaired, and it is unclear how the functional activity of the brain may impact this. Methods: Twenty individual in the chronic stage post-stroke participated in this study. Functional near-infrared spectroscopy (fNIRS) was used to measure prefrontal, pre-motor, sensorimotor, and posterior parietal cortices during walking and walking while completing secondary verbal tasks of varying difficulty. Changes in brain activity during these tasks were measured and relationships were accessed between brain activation changes and cognitive or motor abilities. Results: Significantly larger activations were found for prefrontal, pre-motor, and posterior parietal cortices during dual-task walking. Increasing dual-task walking challenge did not result in an increase in brain activation in these regions. Higher general cognition related to lower increases in activation during the easier dual-task. With the harder dual-task, a trend was also found for higher activation and less motor impairment. Conclusions: This is the first study to show that executive function, motor preparation/planning, and sensorimotor integration areas are all important for dual-task walking post-stroke. A lack of further brain activation increase with increasing challenge suggests a point at which a trade-off between brain activation and performance occurs. Further research is needed to determine if training would result in further increases in brain activity or improved performance.

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Working memory (WM) is a limited-capacity system or set of processes that enables temporary storage and manipulation of information essential for complex cognitive processes. The WM performance is supported by a widespread neural network in which fronto-parietal functional connections have a pivotal role. Transcranial direct current stimulation (tDCS) is rapidly emerging as a promising tool for understanding the role of various cortical areas and their functional networks on cognitive performance. Here we comprehensively evaluated the effects of tDCS on WM by conducting three cross-over counterbalanced sham-controlled experiments in which we contrasted the effects and interactions of the anodal (i.e. facilitatory) tDCS across anterior-posterior (i.e. DLPFC vs PPC) and left-right (i.e. the lateralization) axes, and across online and offline protocols using both verbal and spatial WM (3-back) tasks as outcomes. In the offline protocols, left DLPFC stimulation affected neither verbal nor spatial WM, while left PPC stimulation increased spatial WM. When applied offline over right DLPFC, tDCS improved verbal WM task and marginally enhanced spatial WM; while when tDCS was applied over the right PPC, facilitatory effects were observed on verbal WM. In the online protocol, tDCS did not modulate WM regardless of the task modality or stimulation loci. In summary, the study did not replicate the left DLPFC tDCS effect on WM, found in some of the previous studies, but demonstrated positive effects of stimulation of the right DLPFC as well as PPC bilaterally. The observed effects varied across modality of the 3-back task, and tDCS protocol applied. The results of this study argue for moving towards targeting the lesser-explored stimulation sites within the fronto-parietal network, such as PPC, to gain a better understanding of the usefulness of tDCS for WM neuromodulation.

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Age-related changes in cortico-cortical connectivity in the human motor network in older adults are associated with declines in hand dexterity. Posterior parietal cortex (PPC) is strongly interconnected with motor areas and plays a critical role in many aspects of motor planning. Functional connectivity measures derived from dual-site transcranial magnetic stimulation (dsTMS) studies have found facilitatory inputs from PPC to ipsilateral primary motor cortex (M1) in younger adults. In this study, we investigated whether facilitatory inputs from PPC to M1 are altered by age. We used dsTMS in a conditioning-test paradigm to characterize patterns of functional connectivity between the left PPC and ipsilateral M1 and a standard pegboard test to assess skilled hand motor function in 13 young and 13 older adults. We found a PPC-M1 facilitation in young adults but not older adults. Older adults also showed a decline in motor performance compared to young adults. We conclude that the reduced PPC-M1 facilitation in older adults may be an early marker of age-related decline in the neural control of movement.

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Gait is one of the fundamental behaviors we use to interact with the world. The functionality of the locomotor system is thus related to enriching interactions with our environment. The posterior parietal cortex (PPC) has been found to contribute to motor adaptation during both visuomotor and postural adaptation tasks. Additionally, structural or functional deficits of the PPC lead to impairments in gaits such as shortened steps and increased step width. Based on the aforementioned roles of the PPC, and the importance of gait adaptability, the current investigation sought to identify the role of the PPC in gait adaptation. To achieve this, we performed transcranial direct current stimulation (tDCS) over the bilateral PPC before performing a split-belt treadmill gait adaptation paradigm. We used three stimulation conditions in a within-subject design. tDCS was administered in a randomized and double-blinded order. Following each stimulation session, subjects first performed baseline walking with both belts running at the same speed. Then, subjects walked for 15 min on an uncoupled treadmill, with the belts being driven at a 3:1 speed ratio. Last, they returned to normal (i.e., tied-belt) walking for 5 min. Results from 15 young and healthy subjects identified that subjects required more steps to adapt to split-belt walking following the suppression of the left hemisphere PPC, contralateral to the fast belt. Furthermore, while suppression of the left hemisphere PPC did not increase the number of steps required to re-adapt to tied-belt walking, this condition did lead to increased magnitude of after-effects. Together, these findings indicate that the PPC is involved in locomotor adaptation. These results support previous literature regarding the upper body or postural adaptation and extend these findings to the realm of gait. Results highlight the PPC as a potential target for neurorehabilitation designed to improve gait adaptability.

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Increasing spatial working memory (SWM) load is generally associated with declines in behavioral performance, but the neural correlates of load-related behavioral effects remain poorly understood. Herein, we examine the alterations in oscillatory activity that accompany such performance changes in 22 healthy adults who performed a two- and four-load SWM task during magnetoencephalography (MEG). All MEG data were transformed into the time-frequency domain and significant oscillatory responses were imaged separately per load using a beamformer. Whole-brain correlation maps were computed using the load-related beamformer difference images and load-related accuracy effects on the SWM task. The results indicated that load-related differences in left inferior frontal alpha activity during encoding and maintenance were negatively correlated with load-related accuracy differences on the SWM task. That is, individuals who had more substantial decreases in prefrontal alpha during high-relative to low-load SWM trials tended to have smaller performance decrements on the high-load condition (i.e., they performed more accurately). The same pattern of neurobehavioral correlations was observed during the maintenance period for right superior temporal alpha activity and right superior parietal beta activity. Importantly, this is the first study to employ a voxel-wise whole-brain approach to significantly link load-related oscillatory differences and load-related SWM performance differences.

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Associative memory (AM), an ability to form and retrieve associations between information units is crucial for everyday functioning and is affected by aging as well as by different neurological conditions. It was shown that rTMS over posterior parietal cortex (PPC) can improve AM of face-word pairs. Therefore, we examined if tDCS will produce comparable effects and explore whether the effect would persist one and five days following the stimulation. Thirty-seven healthy participants took part in cross-over sham-controlled study in which they received 20 min of anodal (1.5 mA) or sham tDCS over left PPC. Following tDCS participants completed face-cued word recall and verbal fluency tasks. A randomly selected subsample (N = 18) has completed follow up memory assessments one and five days after the stimulation. Anodal tDCS facilitated AM performance in comparison to sham with the same trend persisting during the 5-day follow-up period. Additionally, participants with lower AM scores had higher relative gain following anodal tDCS. Anodal tDCS had no effect on the control task (verbal fluency). Results support the existence of a specific enhancing effect on AM produced by facilitatory neuromodulation of the PPC. The effect was more prominent in low-performers and it persisted at least 5 days post-stimulation. These findings support the robustness of tDCS effect on AM and provide a foundation for future research that could lead to its future clinical application.

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Associative memory plays a key role in everyday functioning, but it declines with normal ageing as well as due to various pathological states and conditions, thus impairing quality of life. Associative memory enhancement via neurostimulation over frontal areas resulted in limited success, while posterior stimulation sites seemed to be more promising. We hypothesized that anodal transcranial direct current stimulation (tDCS) of parietal areas would lead to higher performance in associative memory due to high connectivity between posterior parietal cortex (PPC) and hippocampus. Forty-two healthy adults participated in two sham-controlled cross-over experiments. Anodal electrode (20 min, 1.5 mA) was placed over P3 in Experiment 1 and over P4 in Experiment 2. During tDCS participants played a simple computer game. After each stimulation session, participants completed parallel forms of an associative memory task (Experiment 1: face-word memory; Experiment 2: object-location memory) and a control task (verbal fluency). In both experiments, associative memory was improved after anodal stimulation compared to sham stimulation, while no differences were observed in the control task. Additionally, memory performance was higher in the second than in the first trial, but the increase in performance between the two trials did not differ between stimulation conditions. It can be concluded that a single-session anodal tDCS over posterior parietal cortex can improve associative memory performance. The specificity, robustness, and reproducibility of the effect suggest that PPC is a promising target for brain stimulation aiming to enhance memory functions.

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BACKGROUND: Time perception associated with durations from 1 s to several minutes involves activity in the right posterior parietal cortex (rPPC). It is unclear whether altering the activity of the rPPC affects an individual's timing performance. Here, we investigated the human timing performance under the application of transcranial direct-current stimulation (tDCS) that altered the neural activities of the rPPC. METHODS: We measured the participants' duration-discrimination threshold by administering a behavioral task during the tDCS application. The tDCS conditions consisted of anodal, cathodal, and sham conditions. The electrodes were placed over the P4 position (10-20 system) and on the left supraorbital forehead. On each task trial, the participant observed two visual stimuli and indicated which was longer. The amount of difference between the two stimulus durations was varied repeatedly throughout the trials according to the participant's responses. The correct answer rate of the trials was calculated for each amount of difference, and the minimum amount with the correct answer rate exceeding 75% was selected as the threshold. The data were analyzed by a linear mixed-effects models procedure. RESULTS: Nineteen volunteers participated in the experiment. We excluded three participants from the analysis: two who reported extreme sleepiness while performing the task and one who could recognize the sham condition correctly with confidence. Our analysis of the 16 participants' data showed that the average value of the thresholds observed under the cathodal condition was lower than that of the sham condition. This suggests that inhibition of the rPPC leads to an improvement in temporal discrimination performance, resulting in improved timing performance. CONCLUSIONS: In the present study, we found a new effect that cathodal tDCS over the rPPC enhances temporal discrimination performance. In terms of the existence of anodal/cathodal tDCS effects on human timing performance, the results were consistent with a previous study that investigated temporal reproduction performance during tDCS application. However, the results of the current study further indicated that cathodal tDCS over the rPPC increases accuracy of observed time duration rather than inducing an overestimation as a previous study reported.

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With a reduced level of alertness, healthy individuals typically show a rightward shift when deploying visual attention in space. The impact of alertness on the neural networks governing visuospatial attention is, however, poorly understood. By using a transcranial magnetic stimulation twin-coil approach, the present study aimed at investigating the effects of an alertness manipulation on the excitability of the left and the right posterior parietal cortices (PPCs), crucial nodes of the visuospatial attentional network. Participants' visuospatial attentional deployment was assessed with a free visual exploration task and concurrent eye tracking. Their alertness level was manipulated through the time of the day, that is, by testing chronotypically defined evening types both during their circadian on- and off-peak times. The results revealed an increased excitability of the left compared with the right PPC during low alertness. On the horizontal dimension, these results were accompanied by a significant rightward shift in the center and a bilateral narrowing in the periphery of the visual exploration field, as well as a central upward shift on the vertical dimension. The findings show that the manipulation of non-spatial attentional aspects (i.e., alertness) can affect visuospatial attentional deployment and modulate the excitability of areas subtending spatial attentional control.

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BACKGROUND: Flexible and precisely timed motor control is based on functional interaction within a cortico-subcortical network. The left posterior parietal cortex (PPC) is supposed to be crucial for anticipatory motor control by sensorimotor feedback matching. OBJECTIVE: Intention of the present study was to disentangle the specific relevance of the left PPC for anticipatory motor control using transcranial direct current stimulation (tDCS) since a causal link remains to be established. METHODS: Anodal vs. cathodal tDCS was applied for 10 min over the left PPC in 16 right-handed subjects in separate sessions. Left primary motor cortex (M1) tDCS served as control condition and was applied in additional 15 subjects. Prior to and immediately after tDCS, subjects performed three tasks demanding temporal motor precision with respect to an auditory stimulus: sensorimotor synchronization as measure of anticipatory motor control, interval reproduction and simple reaction. RESULTS: Left PPC tDCS affected right hand synchronization but not simple reaction times. Motor anticipation was deteriorated by anodal tDCS, while cathodal tDCS yielded the reverse effect. The variability of interval reproduction was increased by anodal left M1 tDCS, whereas it was reduced by cathodal tDCS. No significant effects on simple reaction times were found. CONCLUSION: The present data support the hypothesis that left PPC is causally involved in right hand anticipatory motor control exceeding pure motor implementation as processed by M1 and possibly indicating subjective timing. Since M1 tDCS particularly affects motor implementation, the observed PPC effects are not likely to be explained by alterations of motor-cortical excitability.

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Memories of real and imagined events are qualitatively distinct, and therefore may be supported by different neural mechanisms. In the present study, we tested whether brain regions are differentially activated during source discriminations of perceived versus imagined events. During the encoding phase, subjects perceived and imagined images of objects in response to a cue word. Then, at test, they made judgments about whether old and new cue words corresponded to items that were previously perceived or imagined, or if they were new. The results demonstrated that the left lateral posterior parietal cortex and dorsolateral prefrontal cortex were significantly more active during source attributions of perceived compared to imagined events. In addition, activity in these regions was associated with successful item memory (hits>correct rejections) for perceived, but not imagined events. These findings of a source-based dissociation of successful retrieval activity have important implications regarding theories of parietal contributions to recognition memory.

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