Area-Specificity and Plasticity of History-Dependent Value Coding During Learning.

Decision making is often driven by the subjective value of available options, a value which is formed through experience. To support this fundamental behavior, the brain must encode and maintain the subjective value. To investigate the area specificity and plasticity of value coding, we trained mice in a value-based decision task and imaged neural activity in 6 cortical areas with cellular resolution. History- and value-related signals were widespread across areas, but their strength and temporal patterns differed. In expert mice, the retrosplenial cortex (RSC) uniquely encoded history- and value-related signals with persistent population activity patterns across trials. This unique encoding of RSC emerged during task learning with a strong increase in more distant history signals. Acute inactivation of RSC selectively impaired the reward-history-based behavioral strategy. Our results indicate that RSC flexibly changes its history coding and persistently encodes value-related signals to support adaptive behaviors.

Different roles of response covariability and its attentional modulation in the sensory cortex and posterior parietal cortex.

The covariability of neural responses in the neuron population is highly relevant to the information encoding. Cognitive processes, such as attention, are found to modulate the covariability in the visual cortex to improve information encoding, suggesting the computational advantage of covariability modulation in the neural system. However, is the covariability modulation a general mechanism for enhanced information encoding throughout the information processing pathway, or only adopted in certain processing stages, depending on the property of neural representation? Here, with ultrahigh-field MRI, we examined the covariability, which was estimated by noise correlation, in different attention states in the early visual cortex and posterior parietal cortex (PPC) of the human brain, and its relationship to the quality of information encoding. Our results showed that while attention decreased the covariability to improve the stimulus encoding in the early visual cortex, covariability modulation was not observed in the PPC, where covariability had little impact on information encoding. Further, attention promoted the information flow between the early visual cortex and PPC, with an apparent emphasis on a flow from high- to low-dimensional representations, suggesting the existence of a reduction in the dimensionality of neural representation from the early visual cortex to PPC. Finally, the neural response patterns in the PPC could predict the amplitudes of covariability change in the early visual cortex, indicating a top-down control from the PPC to early visual cortex. Our findings reveal the specific roles of the sensory cortex and PPC during attentional modulation of covariability, determined by the complexity and fidelity of the neural representation in each cortical region.

Mixed Selectivity Coding of Content-Temporal Detail by Dorsomedial Posterior Parietal Neurons.

The dorsomedial posterior parietal cortex (dmPPC) is part of a higher-cognition network implicated in elaborate processes underpinning memory formation, recollection, episode reconstruction, and temporal information processing. Neural coding for complex episodic processing is however under-documented. Here, we recorded extracellular neural activities from three male rhesus macaques (Macaca mulatta) and revealed a set of neural codes of "neuroethogram" in the primate parietal cortex. Analyzing neural responses in macaque dmPPC to naturalistic videos, we discovered several groups of neurons that are sensitive to different categories of ethogram items, low-level sensory features, and saccadic eye movement. We also discovered that the processing of category and feature information by these neurons is sustained by the accumulation of temporal information over a long timescale of up to 30 s, corroborating its reported long temporal receptive windows. We performed an additional behavioral experiment with additional two male rhesus macaques and found that saccade-related activities could not account for the mixed neuronal responses elicited by the video stimuli. We further observed monkeys' scan paths and gaze consistency are modulated by video content. Taken altogether, these neural findings explain how dmPPC weaves fabrics of ongoing experiences together in real time. The high dimensionality of neural representations should motivate us to shift the focus of attention from pure selectivity neurons to mixed selectivity neurons, especially in increasingly complex naturalistic task designs.

Visuomotor prediction during action planning in the human frontoparietal cortex and cerebellum.

The concept of forward models in the brain, classically applied to describing on-line motor control, can in principle be extended to action planning, i.e. assuming forward sensory predictions are issued during the mere preparation of movements. To test this idea, we combined a delayed movement task with a virtual reality based manipulation of visuomotor congruence during functional magnetic resonance imaging. Participants executed simple hand movements after a delay. During the delay, two aspects of the upcoming movement could be cued: the movement type and the visuomotor mapping (i.e. congruence of executed hand movements and visual movement feedback by a glove-controlled virtual hand). Frontoparietal areas showed increased delay period activity when preparing pre-specified movements (cued > uncued). The cerebellum showed increased activity during the preparation for incongruent > congruent visuomotor mappings. The left anterior intraparietal sulcus showed an interaction effect, responding most strongly when a pre-specified (cued) movement was prepared under expected visuomotor incongruence. These results suggest that motor planning entails a forward prediction of visual body movement feedback, which can be adjusted in anticipation of nonstandard visuomotor mappings, and which is likely computed by the cerebellum and integrated with state estimates for (planned) control in the anterior intraparietal sulcus.

Posterior parietal cortex is causally involved in reward valuation but not in probability weighting during risky choice.

This study provides evidence that the posterior parietal cortex is causally involved in risky decision making via the processing of reward values but not reward probabilities. In the within-group experimental design, participants performed a binary lottery choice task following transcranial magnetic stimulation of the right posterior parietal cortex, left posterior parietal cortex, and a right posterior parietal cortex sham (placebo) stimulation. The continuous theta-burst stimulation protocol supposedly downregulating the cortical excitability was used. Both, mean-variance and the prospect theory approach to risky choice showed that the posterior parietal cortex stimulation shifted participants toward greater risk aversion compared with sham. On the behavioral level, after the posterior parietal cortex stimulation, the likelihood of choosing a safer option became more sensitive to the difference in standard deviations between lotteries, compared with sham, indicating greater risk avoidance within the mean-variance framework. We also estimated the shift in prospect theory parameters of risk preferences after posterior parietal cortex stimulation. The hierarchical Bayesian approach showed moderate evidence for a credible change in risk aversion parameter toward lower marginal reward value (and, hence, lower risk tolerance), while no credible change in probability weighting was observed. In addition, we observed anecdotal evidence for a credible increase in the consistency of responses after the left posterior parietal cortex stimulation compared with sham.

Resting-state functional connectivity involved in tactile orientation processing.

BACKGROUND: Grating orientation discrimination (GOD) is commonly used to assess somatosensory spatial processing. It allows discrimination between parallel and orthogonal orientations of tactile stimuli applied to the fingertip. Despite its widespread application, the underlying mechanisms of GOD, particularly the role of cortico-cortical interactions and local brain activity in this process, remain elusive. Therefore, we aimed to investigate how a specific cortico-cortical network and inhibitory circuits within the primary somatosensory cortex (S1) and secondary somatosensory cortex (S2) contribute to GOD. METHODS: In total, 51 healthy young adults were included in our study. We recorded resting-state magnetoencephalography (MEG) and somatosensory-evoked magnetic field (SEF) in participants with open eyes. We converted the data into a source space based on individual structural magnetic resonance imaging. Next, we estimated S1- and S2-seed resting-state functional connectivity (rs-FC) at the alpha and beta bands through resting-state MEG using the amplitude envelope correlation method across the entire brain (i.e., S1/S2-seeds × 15,000 vertices × two frequencies). We assessed the inhibitory response in the S1 and S2 from SEFs using a paired-pulse paradigm. We automatically measured the GOD task in parallel and orthogonal orientations to the index finger, applying various groove widths with a custom-made device. RESULTS: We observed a specific association between the GOD threshold (all P < 0.048) and the alpha rs-FC in the S1-superior parietal lobule and S1-adjacent to the parieto-occipital sulcus (i.e., lower rs-FC values corresponded to higher performance). In contrast, no association was observed between the local responses and the threshold. DISCUSSION: The results of this study underpin the significance of specific cortico-cortical networks in recognizing variations in tactile stimuli.

Disentangling Cerebellar and Parietal Contributions to Gait and Body Schema: A Repetitive Transcranial Magnetic Stimulation Study.

The overlap between motor and cognitive signs resulting from posterior parietal cortex (PPC) and cerebellar lesions can mask their relative contribution in the sensorimotor integration process. This study aimed to identify distinguishing motor and cognitive features to disentangle PPC and cerebellar involvement in two sensorimotor-related functions: gait and body schema representation. Thirty healthy volunteers were enrolled and randomly assigned to PPC or cerebellar stimulation. Sham stimulation and 1 Hz-repetitive-Transcranial-Magnetic-Stimulation were delivered over P3 or cerebellum before a balance and a walking distance estimation task. Each trial was repeated with eyes open (EO) and closed (EC). Eight inertial measurement units recorded spatiotemporal and kinematic variables of gait. Instability increased in both groups after real stimulation: PPC inhibition resulted in increased instability in EC conditions, as evidenced by increased ellipse area and range of movement in medio-lateral and anterior-posterior (ROMap) directions. Cerebellar inhibition affected both EC (increased ROMap) and EO stability (greater displacement of the center of mass). Inhibitory stimulation (EC vs. EO) affected also gait spatiotemporal variability, with a high variability of ankle and knee angles plus different patterns in the two groups (cerebellar vs parietal). Lastly, PPC group overestimates distances after real stimulation (EC condition) compared to the cerebellar group. Stability, gait variability, and distance estimation parameters may be useful clinical parameters to disentangle cerebellar and PPC sensorimotor integration deficits. Clinical differential diagnosis efficiency can benefit from this methodological approach.

Complementary contribution of the medial and lateral human parietal cortex to grasping: a repetitive TMS study.

The dexterous control of our grasping actions relies on the cooperative activation of many brain areas. In the parietal lobe, 2 grasp-related areas collaborate to orchestrate an accurate grasping action: dorsolateral area AIP and dorsomedial area V6A. Single-cell recordings in monkeys and fMRI studies in humans have suggested that both these areas specify grip aperture and wrist orientation, but encode these grasping parameters differently, depending on the context. To elucidate the causal role of phAIP and hV6A, we stimulated these areas, while participants were performing grasping actions (unperturbed grasping). rTMS over phAIP impaired the wrist orientation process, whereas stimulation over hV6A impaired grip aperture encoding. In a small percentage of trials, an unexpected reprogramming of grip aperture or wrist orientation was required (perturbed grasping). In these cases, rTMS over hV6A or over phAIP impaired reprogramming of both grip aperture and wrist orientation. These results represent the first direct demonstration of a different encoding of grasping parameters by 2 grasp-related parietal areas.

Sensory modality and spoken language shape reading network in blind readers of Braille.

The neural basis of reading is highly consistent across many languages and scripts. Are there alternative neural routes to reading? How does the sensory modality of symbols (tactile vs. visual) influence their neural representations? We examined these questions by comparing reading of visual print (sighted group, n = 19) and tactile Braille (congenitally blind group, n = 19). Blind and sighted readers were presented with written (words, consonant strings, non-letter shapes) and spoken stimuli (words, backward speech) that varied in word-likeness. Consistent with prior work, the ventral occipitotemporal cortex (vOTC) was active during Braille and visual reading. A posterior/anterior vOTC word-form gradient was observed only in sighted readers with more anterior regions preferring larger orthographic units (words). No such gradient was observed in blind readers. Consistent with connectivity predictions, in blind compared to sighted readers, posterior parietal cortices were recruited to a greater degree and contained word-preferring patches. Lateralization of Braille in blind readers was predicted by laterality of spoken language and reading hand. The effect of spoken language increased along a cortical hierarchy, whereas effect of reading hand waned. These results suggested that the neural basis of reading is influenced by symbol modality and spoken language and support connectivity-based views of cortical function.

More Than the Sum of Its Parts: Visual-Tactile Integration in the Behaving Rat.

We experience the world by constantly integrating cues from multiple modalities to form unified sensory percepts. Once familiar with multimodal properties of an object, we can recognize it regardless of the modality involved. In this chapter we will examine the case of a visual-tactile orientation categorization experiment in rats. We will explore the involvement of the cerebral cortex in recognizing objects through multiple sensory modalities. In the orientation categorization task, rats learned to examine and judge the orientation of a raised, black and white grating using touch, vision, or both. Their multisensory performance was better than the predictions of linear models for cue combination, indicating synergy between the two sensory channels. Neural recordings made from a candidate associative cortical area, the posterior parietal cortex (PPC), reflected the principal neuronal correlates of the behavioral results: PPC neurons encoded both graded information about the object and categorical information about the animal's decision. Intriguingly single neurons showed identical responses under each of the three modality conditions providing a substrate for a neural circuit in the cortex that is involved in modality-invariant processing of objects.

Contralateral Limb Specificity for Movement Preparation in the Parietal Reach Region.

The canonical view of motor control is that distal musculature is controlled primarily by the contralateral cerebral hemisphere; unilateral brain lesions typically affect contralateral but not ipsilateral musculature. Contralateral-only limb deficits following a unilateral lesion suggest but do not prove that control is strictly contralateral: the loss of a contribution of the lesioned hemisphere to the control of the ipsilesional limb could be masked by the intact contralateral drive from the nonlesioned hemisphere. To distinguish between these possibilities, we serially inactivated the parietal reach region, comprising the posterior portion of medial intraparietal area, the anterior portion of V6a, and portions of the lateral occipital parietal area, in each hemisphere of 2 monkeys (23 experimental sessions, 46 injections total) to evaluate parietal reach region's contribution to the contralateral reaching deficits observed following lateralized brain lesions. Following unilateral inactivation, reach reaction times with the contralesional limb were slowed compared with matched blocks of control behavioral data; there was no effect of unilateral inactivation on the reaction time of either ipsilesional limb reaches or saccadic eye movements. Following bilateral inactivation, reaching was slowed in both limbs, with an effect size in each no different from that produced by unilateral inactivation. These findings indicate contralateral organization of reach preparation in posterior parietal cortex.SIGNIFICANCE STATEMENT Unilateral brain lesions typically affect contralateral but not ipsilateral musculature. Contralateral-only limb deficits following a unilateral lesion suggest but do not prove that control is strictly contralateral: the loss of a contribution of the lesioned hemisphere to the control of the ipsilesional limb could be masked by the intact contralateral drive from the nonlesioned hemisphere. Unilateral lesions cannot distinguish between contralateral and bilateral control, but bilateral lesions can. Here we show similar movement initiation deficits after combined unilateral and bilateral inactivation of the parietal reach region, indicating contralateral organization of reach preparation.

Dissociation between Attention-Dependent and Spatially Specific Illusory Shape Responses within the Topographic Areas of the Posterior Parietal Cortex.

The human visual system consists of multiple topographic maps that extend from the early visual cortex (EVC) along the dorsal and ventral processing streams. Responses to illusory shapes within these maps have been demonstrated in the ventral stream areas, in particular the lateral occipital complex (LOC). Recently, the intraparietal sulcus (IPS) of the dorsal stream has been linked to the processing of illusory shapes defined by motion. It remains unclear whether the topographically organized parietal areas also respond to stationary illusory shapes, which would suggest their generic role in representing illusory content. In the current study we measured brain responses using fMRI while 30 human participants (12 male) observed flickering inducers around the fixation task. The inducers either formed an illusory diamond in the center, a triangle in the left or right hemifield, or were inverted such that no illusory figure was formed. We compared responses of parietal regions IPS0-IPS5 and SPL1 to each illusory figure with the nonillusory condition. To determine the role of attentional modulation on illusory shape responses we manipulated the difficulty of the fixation task. Our results show that all IPS areas responded to illusory shapes. The more posterior areas IPS0-IPS3 additionally displayed a preference toward contralateral shapes, while the more anterior areas IPS4 and IPS5 showed response attenuation with increased task difficulty. We suggest that the IPS can represent illusory content generated not only by moving, but also by stationary stimuli, and that there is a functional dissociation between attention-dependent anterior and spatially specific posterior topographic maps.SIGNIFICANCE STATEMENT The traditional view of the ventral visual pathway being solely responsible for representation of objects has recently been challenged by demonstrating illusory shape representation within the dorsal visual pathway with moving bistable stimuli. Our results provide evidence for the dorsal stream contribution to representing not only moving, but also stationary illusory shapes. Our results also show a functional subdivision along the topographic maps, with spatially specific shape responses in the more posterior, and attention-dependent responses in the more anterior areas. These findings have implications for our understanding of the relationship between attention and grouping in healthy individuals and neuropsychological patients. Furthermore, IPS areas should be considered in theoretical accounts and models of how subjective content is generated in the brain.

The Neural Bases of Action-Outcome Learning in Humans.

From an associative perspective the acquisition of new goal-directed actions requires the encoding of specific action-outcome (AO) associations and, therefore, sensitivity to the validity of an action as a predictor of a specific outcome relative to other events. Although competitive architectures have been proposed within associative learning theory to achieve this kind of identity-based selection, whether and how these architectures are implemented by the brain is still a matter of conjecture. To investigate this issue, we trained human participants to encode various AO associations while undergoing functional neuroimaging (fMRI). We then degraded one AO contingency by increasing the probability of the outcome in the absence of its associated action while keeping other AO contingencies intact. We found that this treatment selectively reduced performance of the degraded action. Furthermore, when a signal predicted the unpaired outcome, performance of the action was restored, suggesting that the degradation effect reflects competition between the action and the context for prediction of the specific outcome. We used a Kalman filter to model the contribution of different causal variables to AO learning and found that activity in the medial prefrontal cortex (mPFC) and the dorsal anterior cingulate cortex (dACC) tracked changes in the association of the action and context, respectively, with regard to the specific outcome. Furthermore, we found the mPFC participated in a network with the striatum and posterior parietal cortex to segregate the influence of the various competing predictors to establish specific AO associations.SIGNIFICANCE STATEMENT Humans and other animals learn the consequences of their actions, allowing them to control their environment in a goal-directed manner. Nevertheless, it is unknown how we parse environmental causes from the effects of our own actions to establish these specific action-outcome (AO) relationships. Here, we show that the brain learns the causal structure of the environment by segregating the unique influence of actions from other causes in the medial prefrontal and anterior cingulate cortices and, through a network of structures, including the caudate nucleus and posterior parietal cortex, establishes the distinct causal relationships from which specific AO associations are formed.

Respiratory entrainment of units in the mouse parietal cortex depends on vigilance state.

Synchronous oscillations are essential for coordinated activity in neuronal networks and, hence, for behavior and cognition. While most network oscillations are generated within the central nervous system, recent evidence shows that rhythmic body processes strongly influence activity patterns throughout the brain. A major factor is respiration (Resp), which entrains multiple brain regions at the mesoscopic (local field potential) and single-cell levels. However, it is largely unknown how such Resp-driven rhythms interact or compete with internal brain oscillations, especially those with similar frequency domains. In mice, Resp and theta (θ) oscillations have overlapping frequencies and co-occur in various brain regions. Here, we investigated the effects of Resp and θ on neuronal discharges in the mouse parietal cortex during four behavioral states which either show prominent θ (REM sleep and active waking (AW)) or lack significant θ (NREM sleep and waking immobility (WI)). We report a pronounced state-dependence of spike modulation by both rhythms. During REM sleep, θ effects on unit discharges dominate, while during AW, Resp has a larger influence, despite the concomitant presence of θ oscillations. In most states, unit modulation by θ or Resp increases with mean firing rate. The preferred timing of Resp-entrained discharges (inspiration versus expiration) varies between states, indicating state-specific and different underlying mechanisms. Our findings show that neurons in an associative cortex area are differentially and state-dependently modulated by two fundamentally different processes: brain-endogenous θ oscillations and rhythmic somatic feedback signals from Resp.

Single-neuron and population measures of neuronal activity in working memory tasks.

Information represented in working memory is reflected in the firing rate of neurons in the prefrontal cortex and brain areas connected to it. In recent years, there has been an increased realization that population measures capture more accurately neural correlates of cognitive functions. We examined how single neuron firing in the prefrontal and posterior parietal cortex of two male monkeys compared with population measures in spatial working memory tasks. Persistent activity was observed in the dorsolateral prefrontal and posterior parietal cortex and firing rate predicted working memory behavior, particularly in the prefrontal cortex. These findings had equivalents in population measures, including trajectories in state space that became less separated in error trials. We additionally observed rotations of stimulus representations in the neuronal state space for different task conditions, which were not obvious in firing rate measures. These results suggest that population measures provide a richer view of how neuronal activity is associated with behavior, largely confirming that persistent activity is the core phenomenon that maintains visual-spatial information in working memory.NEW & NOTEWORTHY Recordings from large numbers of neurons led to a reevaluation of neural correlates of cognitive functions, which traditionally were defined based on responses of single neurons or averages of firing rates. Analysis of neuronal recordings from the dorsolateral prefrontal and posterior parietal cortex revealed that properties of neuronal firing captured in classical studies of persistent activity can account for population representations, though some population characteristics did not have clear correlates in single neuron activity.

Direction of visual shift and hand congruency enhance spatial realignment during visuomotor adaptation.

Although prism adaptation has been studied extensively for over 100 years to better understand how the motor system adapts to sensory perturbations, very few studies have systematically studied how the combination of the hand used to adapt, and the direction of visual shift, might influence adaptation. Given that sensory inputs and motor outputs from the same side are processed (at least initially) in the same hemisphere, we wondered whether there might be differences in how people adapt when the hand used and the direction of visual shift were congruent (e.g., adapting to rightward shifting prisms with the right hand), compared to incongruent (e.g., adapting to rightward shifting prisms with the left hand). In Experiment 1 we re-analyzed a previously published dataset (Striemer, Enns, and Whitwell Striemer et al., Cortex 115:201-215, 2019a) in which healthy adults (n = 17) adapted to 17° leftward or rightward optically displacing prisms using their left or right hand (tested in separate sessions, counterbalanced). Our results revealed a "congruency effect" such that adaptation aftereffects were significantly larger for reaches performed without visual feedback (i.e., straight-ahead pointing) when the direction of prism shift and the hand used were congruent, compared to incongruent. We replicated this same congruency effect in Experiment 2 in a new group of participants (n = 25). We suggest that a better understanding of the cognitive and neural mechanisms underlying the congruency effect will allow researchers to build more precise models of visuomotor learning, and may lead to the development of more effective applications of prism adaptation for the treatment of attentional disorders following brain damage.

Neural Basis of Somatosensory Spatial and Temporal Discrimination in Humans: The Role of Sensory Detection.

While detecting somatic stimuli from the external environment, an accurate determination of their spatial and temporal properties is essential for human behavior. Whether and how detection relates to human capacity for somatosensory spatial discrimination (SD) and temporal discrimination (TD) remains unclear. Here, participants underwent functional magnetic resonance imaging scanning when simply detecting vibrotactile stimuli of the leg, judging their location (SD), or deciding their number in time (TD). By conceptualizing tactile discrimination as consisting of detection and determination processes, we found that tactile detection elicited activation specifically involved in SD within the right inferior and superior parietal lobules, 2 regions previously implicated in the control of spatial attention. These 2 regions remained activated in the determination process, during which functional connectivity between these 2 regions predicted individual SD ability. In contrast, tactile detection produced little activation specifically related to TD. Participants' TD ability was implemented in brain regions implicated in coding temporal structures of somatic stimuli (primary somatosensory cortex) and time estimation (anterior cingulate, pre-supplementary motor area, and putamen). Together, our findings indicate a close link between somatosensory detection and SD (but not TD) at the neural level, which aids in explaining why we can promptly respond toward detected somatic stimuli.

Exploring Cognition with Brain-Machine Interfaces.

Traditional brain-machine interfaces decode cortical motor commands to control external devices. These commands are the product of higher-level cognitive processes, occurring across a network of brain areas, that integrate sensory information, plan upcoming motor actions, and monitor ongoing movements. We review cognitive signals recently discovered in the human posterior parietal cortex during neuroprosthetic clinical trials. These signals are consistent with small regions of cortex having a diverse role in cognitive aspects of movement control and body monitoring, including sensorimotor integration, planning, trajectory representation, somatosensation, action semantics, learning, and decision making. These variables are encoded within the same population of cells using structured representations that bind related sensory and motor variables, an architecture termed partially mixed selectivity. Diverse cognitive signals provide complementary information to traditional motor commands to enable more natural and intuitive control of external devices.

Mechanistic determinants of effector-independent motor memory encoding.

Coordinated, purposeful movements learned with one effector generalize to another effector, a finding that has important implications for tool use, sports, performing arts, and rehabilitation. This occurs because the motor memory acquired through learning comprises representations that are effector-independent. Despite knowing this for decades, the neural mechanisms and substrates that are causally associated with the encoding of effector-independent motor memories remain poorly understood. Here we exploit intereffector generalization, the behavioral signature of effector-independent representations, to address this crucial gap. We first show in healthy human participants that postlearning generalization across effectors is principally predicted by the level of an implicit mechanism that evolves gradually during learning to produce a temporally stable memory. We then demonstrate that interfering with left but not right posterior parietal cortex (PPC) using high-definition cathodal transcranial direct current stimulation impedes learning mediated by this mechanism, thus potentially preventing the encoding of effector-independent memory components. We confirm this in our final experiment in which we show that disrupting left PPC but not primary motor cortex after learning has been allowed to occur blocks intereffector generalization. Collectively, our results reveal the key mechanism that encodes an effector-independent memory trace and uncover a central role for the PPC in its representation. The encoding of such motor memory components outside primary sensorimotor regions likely underlies a parsimonious neural organization that enables more efficient movement planning in the brain, independent of the effector used to act.

Brain activity patterns underlying memory confidence.

The primary aim of this review is to examine the brain activity patterns that are related to subjectively perceived memory confidence. We focus on the main brain regions involved in episodic memory: the medial temporal lobe (MTL), prefrontal cortex (PFC), and posterior parietal cortex (PPC), and relate activity in their subregions to memory confidence. How this brain activity in both the encoding and retrieval phase is related to (subsequent) memory confidence ratings will be discussed. Specifically, encoding related activity in MTL regions and ventrolateral PFC mainly shows a positive linear increase with subsequent memory confidence, while dorsolateral and ventromedial PFC activity show mixed patterns. In addition, encoding-related PPC activity seems to only have indirect effects on memory confidence ratings. Activity during retrieval in both the hippocampus and parahippocampal cortex increases with memory confidence, especially during high-confident recognition. Retrieval-related activity in the PFC and PPC show mixed relationships with memory confidence, likely related to post-retrieval monitoring and attentional processes, respectively. In this review, these MTL, PFC, and PPC activity patterns are examined in detail and related to their functional roles in memory processes. This insight into brain activity that underlies memory confidence is important for our understanding of brain-behaviour relations and memory-guided decision making.