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Dorsal pulvinar has been implicated in visuospatial attentional and perceptual confidence processing. Perturbations of the dorsal pulvinar also induce an overt spatial saccade bias during free choices. But it remains unclear whether the dorsal pulvinar inactivation during an oculomotor target selection based on a perceptual decision will lead to perceptual impairment or a more general orienting deficit. To address this question, we reversibly inactivated unilateral dorsal pulvinar by injecting GABA-A agonist THIP while two macaque monkeys performed a color discrimination saccade response task with varying perceptual difficulty. We used Signal Detection Theory to dissociate perceptual discrimination (dprime) and spatial selection bias (response criterion) effects. We expected a decrease in dprime if dorsal pulvinar affects perceptual discrimination and a shift in response criterion if dorsal pulvinar is mainly involved in spatial orienting. After inactivation, we observed response criterion shifts away from contralesional stimuli, especially when two competing peripheral stimuli in opposite hemifields were present, for both difficulty levels. The saccade latency for the contralesional selection increased under all conditions. Notably, the dprime and overall accuracy remained largely unaffected. Our results underline the critical contribution of the dorsal pulvinar to spatial orienting while being less important for perceptual discrimination.

Standard neuroeconomics theories state that the value of different classes of stimuli, for instance the hedonic value of food versus music, is transformed to a common reference scale that is independent of their sensory properties. However, adaptive behaviour in a multimodal and dynamic environment requires that our brain also encodes information about the sensory features of reward predicting stimuli. Whether and how a common code for value could integrate information about the sensory features of rewarding stimuli remains inadequately understood. By employing stimuli from auditory and visual modalities as reward predicting cues in a value-based decision-making task, we were able to vary the reward value and sensory modality independently and dissociate neural codes of auditory and visual rewards in frontal areas using fMRI. Univariate fMRI analysis revealed modality-specific and modality-general value representations in orbitofrontal cortex (OFC) and ventromedial prefrontal cortex (vmPFC), respectively. Crucially, modality-specific representations were highly selective as they were only activated when participants believed that the corresponding sensory modality was associated with reward and were absent when the task involved instruction-based rather than value-based choices. Moreover, we show that modality-specific value representations are supported by the presence of the effective connectivity between each primary sensory area and the corresponding OFC activation and further between modality-specific value representations in OFC and vmPFC, only when the sensory modality to be chosen is associated with reward and absent otherwise. Our results indicate the presence of both modality-specific and modality-general representations of reward value and reveal mechanisms through which the interaction between the sensory cortices and the two types of representation guides value-based decisions.

When comparing themselves with others, people often perceive their own actions and behaviour favourably. This phenomenon is often categorised as a bias of attribution, with favourable self-evaluation resulting from differing explanations of one’s own behaviour and that of others. However, studies on availability biases offer an alternative explanation, ascribing egocentric biases to the inherent sensory asymmetries between performing an action and merely observing it. In this study, we used a paradigm that allowed us to directly compare these two distinct sources of bias. Participants perceived the tasks they performed to be harder than the tasks they observed, but demonstrated no bias driven by favourable self-evaluation. Furthermore, the degree of overestimation of the difficulty of performed tasks was magnified as overall task difficulty increased. These findings suggest that egocentric biases are in part derived from sensory asymmetries inherent to the first-person perspective.

Visual perceptual learning is traditionally thought to arise in visual cortex. However, typical perceptual learning tasks also involve systematic mapping of visual information onto motor actions. Because the motor system contains both effector-specific and effector-unspecific representations, the question arises whether visual perceptual learning is effector-specific itself, or not. Here, we study this question in an orientation discrimination task. Subjects learn to indicate their choices either with joystick movements or with manual reaches. After training, we challenge them to perform the same task with eye movements. We dissect the decision-making process using the drift diffusion model. We find that learning effects on the rate of evidence accumulation and the decision criteria depend on effectors, albeit not fully. This suggests that during perceptual learning, visual information is mapped onto effector-specific integrators. Overlap of the populations of neurons encoding motor plans for these effectors may explain partial generalization. Taken together, visual perceptual learning is not limited to visual cortex, but also affects sensorimotor mapping at the interface of visual processing and decision making.

Contains fulltext : 239574.pdf (Publisher’s version ) (Closed access) Open science initiatives are creating opportunities to increase research coordination and impact in nonhuman primate (NHP) imaging. The PRIMatE Data and Resource Exchange community recently developed a collaboration-based strategic plan to advance NHP imaging as an integrative approach for multiscale neuroscience. 5 p.

Causal perturbations suggest that the primate dorsal pulvinar (dPul) plays a crucial role in target selection and saccade planning, but its basic visuomotor neuronal properties are unclear. While some functional aspects of dPul and interconnected frontoparietal areas – e.g. ipsilesional choice bias after inactivation – are similar, it is unknown if dPul shares oculomotor properties of the cortical circuitry, in particular the delay and choice-related activity. We investigated such properties in macaque dPul during instructed and free-choice memory saccades. Most recorded units showed visual (16%), visuomotor (29%) or motor-related (35%) responses. Visual responses were mainly contralateral; motor-related responses were predominantly post-saccadic (64%) and showed weak contralateral bias. Pre-saccadic enhancement was infrequent (9-15%) – instead, activity was often suppressed during saccade planning (30%) and execution (19%). Surprisingly, only few units exhibited classical visuomotor patterns combining cue and continuous delay activity until the saccade or pre-saccadic ramping, and most spatially-selective neurons did not encode the upcoming decision during free-choice delay. Thus, in absence of a visible goal, the dorsal pulvinar has a limited role in prospective saccade planning, with patterns partially complementing its frontoparietal partners. Conversely, prevalent cue and post-saccadic responses imply the participation in integrating spatial goals with processing across saccades.

Estimating invested effort is a core dimension for evaluating own and others’ actions, and views on the relationship between effort and rewards are deeply ingrained in various societal attitudes. Internal representations of effort, however, are inherently noisy, e.g. due to the variability of sensorimotor and visceral responses to physical exertion. The uncertainty in effort judgments is further aggravated when there is no direct access to the internal representations of exertion – such as when estimating the effort of another person. Bayesian cue integration suggests that this uncertainty can be resolved by incorporating additional cues that are predictive of effort, e.g. received rewards. We hypothesized that judgments about the effort spent on a task will be influenced by the magnitude of received rewards. Additionally, we surmised that such influence might further depend on individual beliefs regarding the relationship between hard work and prosperity, as exemplified by a conservative work ethic. To test these predictions, participants performed an effortful task interleaved with a partner and were informed about the obtained reward before rating either their own or the partner’s effort. We show that higher rewards led to higher estimations of exerted effort in self-judgments, and this effect was even more pronounced for other-judgments. In both types of judgment, computational modelling revealed that reward information and sensorimotor markers of exertion were combined in a Bayes-optimal manner in order to reduce uncertainty. Remarkably, the extent to which rewards influenced effort judgments was associated with conservative world-views, indicating links between this phenomenon and general beliefs about the relationship between effort and earnings in society. Open-Access-Publikationsfonds 2020 peerReviewed

Despite many years of intense research, there is no strong consensus about the role of the lateral intraparietal area (LIP) in decision making. One view of LIP function is that it guides spatial attention, providing a “saliency map” of the external world. If this were the case, it would contribute to target selection regardless of which action would be performed to implement the choice. On the other hand, LIP inactivation has been shown to influence spatial selection and oculomotor metrics in free-choice decisions, which are made using eye movements, arguing that it contributes to saccade decisions. To dissociate between a more general attention role and a more effector specific saccade role, we reversibly inactivated LIP while non-human primates freely selected between two targets, presented in the two hemifields, with either saccades or reaches. Unilateral LIP inactivation induced a strong choice bias to ipsilesional targets when decisions were made with saccades. Interestingly, the inactivation also caused a reduction of contralesional choices when decisions were made with reaches, albeit the effect was less pronounced. These findings suggest that LIP is part of a network for making oculomotor decisions and is largely effector-specific in free-choice decisions.

Expansion of the dorsal pulvinar in humans and its anatomical connectivity suggests its involvement in higher-order cognitive and visuomotor functions. We investigated visuomotor performance in a 31 year old patient (M.B.) with a lesion centered on the medial portion of the dorsal pulvinar (left > right) due to an atypical Sarcoidosis manifestation. Unlike lesions with a vascular etiology, the lesion of M.B. did not include primary sensory or motor thalamic nuclei. Thus, this patient gave us the exceedingly rare opportunity to study the contribution of the dorsal pulvinar to visuomotor behavior in a human without confounding losses in primary sensory or motor domains. We investigated reaching, saccade and visual decision making performance. Patient data in each task was compared to at least seven age matched healthy controls. While saccades were hypometric towards both hemifields, the patient did not show any spatial choice bias or perceptual deficits. At the same time, he exhibited reach and grasp difficulties, which shared features with both, parietal and cerebellar damage. In particular, he had problems to form a precision grip and exhibited reach deficits expressed in decreased accuracy, delayed initiation and prolonged movement durations. Reach deficits were similar in foveal and extrafoveal viewing conditions and in both visual hemifields but were stronger with the right hand. These results suggest that dorsal pulvinar function in humans goes beyond its subscribed role in visual cognition and is critical for the programming of voluntary actions with the hands.

Much evidence indicates that humans and other species process large-scale visual information before fine spatial detail. Neurophysiological data obtained with paralyzed eyes suggest that this coarse-to-fine sequence results from spatiotemporal filtering by neurons in the early visual pathway. However, the eyes are normally never stationary: rapid gaze shifts (saccades) incessantly alternate with slow fixational movements. To investigate the consequences of this oculomotor cycle on the dynamics of perception, we combined spectral analysis of visual input signals, neural modeling, and gaze-contingent control of retinal stimulation in humans. We show that the saccade/fixation cycle reformats the flow impinging on the retina in a way that initiates coarse-to-fine processing at each fixation. This finding reveals that the visual system uses oculomotor-induced temporal modulations to sequentially encode different spatial components and suggests that, rather than initiating coarse-to-fine processing, spatiotemporal coupling in the early visual pathway builds on the information dynamics of the oculomotor cycle.

Contains fulltext : 217200.pdf (Publisher’s version ) (Closed access) Nonhuman primate neuroimaging is on the cusp of a transformation, much in the same way its human counterpart was in 2010, when the Human Connectome Project was launched to accelerate progress. Inspired by an open data-sharing initiative, the global community recently met and, in this article, breaks through obstacles to define its ambitions. 4 p.

Sensorimotor cortical areas contain eye position information thought to ensure perceptual stability across saccades and underlie spatial transformations supporting goal-directed actions. One pathway by which eye position signals could be relayed to and across cortical areas is via the dorsal pulvinar. Several studies have demonstrated saccade-related activity in the dorsal pulvinar, and we have recently shown that many neurons exhibit postsaccadic spatial preference. In addition, dorsal pulvinar lesions lead to gaze-holding deficits expressed as nystagmus or ipsilesional gaze bias, prompting us to investigate the effects of eye position. We tested three starting eye positions (−15°, 0°, 15°) in monkeys performing a visually cued memory saccade task. We found two main types of gaze dependence. First, ~50% of neurons showed dependence on static gaze direction during initial and postsaccadic fixation, and might be signaling the position of the eyes in the orbit or coding foveal targets in a head/body/world-centered reference frame. The population-derived eye position signal lagged behind the saccade. Second, many neurons showed a combination of eye-centered and gaze-dependent modulation of visual, memory, and saccadic responses to a peripheral target. A small subset showed effects consistent with eye position-dependent gain modulation. Analysis of reference frames across task epochs from visual cue to postsaccadic fixation indicated a transition from predominantly eye-centered encoding to representation of final gaze or foveated locations in nonretinocentric coordinates. These results show that dorsal pulvinar neurons carry information about eye position, which could contribute to steady gaze during postural changes and to reference frame transformations for visually guided eye and limb movements. NEW & NOTEWORTHY Work on the pulvinar focused on eye-centered visuospatial representations, but position of the eyes in the orbit is also an important factor that needs to be taken into account during spatial orienting and goal-directed reaching. We show that dorsal pulvinar neurons are influenced by eye position. Gaze direction modulated ongoing firing during stable fixation, as well as visual and saccade responses to peripheral targets, suggesting involvement of the dorsal pulvinar in spatial coordinate transformations.

Following the expanding use and applications of virtual reality in every-day life, dynamic virtual stimuli are of increasing interest in cognitive studies. They allow for control of features such as gaze, expression and movement, which may help to overcome limitations of using either static or poorly controlled real stimuli. In using virtual stimuli however, one must be careful to avoid the uncanny valley effect - where realistic stimuli can be perceived as eerie, and induce an aversion response. At the same time, it is important to establish whether responses to virtual stimuli mirror responses to depictions of a real conspecific. In the current study, we describe the development of a new avatar with realistic features for nonhuman primates, the ‘primatar’. As a first step towards validation, we assessed how monkeys respond to images of this avatar compared to images of real monkeys, and an unrealistic avatar. We also compared responses between original images and scrambled as well as obfuscated versions of these images. We measured looking time to images in six free moving long-tailed macaques (Macaca fascicularis) and eye movement exploration behaviour in three rhesus macaques (Macaca mulatta). Both groups showed more of such signs of overt attention to original images than scrambled or obfuscated images. In addition, we assessed whether the realistic avatar created an uncanny valley effect through decreased looking time, finding that in both groups, monkeys did not differentiate between real, realistic or unrealistic images. These results provide support for further development of our avatar for use in social cognition studies, and more generally for cognitive research with virtual stimuli in nonhuman primates. Future research needs to shed light on the source of the inconsistent findings for the uncanny valley effect in macaques, to elucidate the roots of this mechanism in humans.

The pulvinar complex is interconnected extensively with brain regions involved in spatial processing and eye movement control. Recent inactivation studies have shown that the dorsal pulvinar (dPul) plays a role in saccade target selection; however, it remains unknown whether it exerts effects on visual processing or at planning/execution stages. We used electrical microstimulation of the dPul while monkeys performed saccade tasks toward instructed and freely chosen targets. Timing of stimulation was varied, starting before, at, or after onset of target(s). Stimulation affected saccade properties and target selection in a time-dependent manner. Stimulation starting before but overlapping with target onset shortened saccadic reaction times (RTs) for ipsiversive (to the stimulation site) target locations, whereas stimulation starting at and after target onset caused systematic delays for both ipsiversive and contraversive locations. Similarly, stimulation starting before the onset of bilateral targets increased ipsiversive target choices, whereas stimulation after target onset increased contraversive choices. Properties of dPul neurons and stimulation effects were consistent with an overall contraversive drive, with varying outcomes contingent upon behavioral demands. RT and choice effects were largely congruent in the visually-guided task, but stimulation during memory-guided saccades, while influencing RTs and errors, did not affect choice behavior. Together, these results show that the dPul plays a primary role in action planning as opposed to visual processing, that it exerts its strongest influence on spatial choices when decision and action are temporally close, and that this choice effect can be dissociated from motor effects on saccade initiation and execution.SIGNIFICANCE STATEMENTDespite a recent surge of interest, the core function of the pulvinar, the largest thalamic complex in primates, remains elusive. This understanding is crucial given the central role of the pulvinar in current theories of integrative brain functions supporting cognition and goal-directed behaviors, but electrophysiological and causal interference studies of dorsal pulvinar (dPul) are rare. Building on our previous studies that pharmacologically suppressed dPul activity for several hours, here we used transient electrical microstimulation at different periods while monkeys performed instructed and choice eye movement tasks, to determine time-specific contributions of pulvinar to saccade generation and decision making. We show that stimulation effects depend on timing and behavioral state and that effects on choices can be dissociated from motor effects.

A Transparent game is a game-theoretic setting that takes action visibility into account. In each round, depending on the relative timing of their actions, players have a certain probability to see their partner’s choice before making their own decision. This probability is determined by the level of transparency. At the two extremes, a game with zero transparency is equivalent to the classical simultaneous game, and a game with maximal transparency corresponds to a sequential game. Despite the prevalence of intermediate transparency in many everyday interactions such scenarios have not been sufficiently studied. Here we consider a transparent iterated Prisoner’s dilemma (iPD) and use evolutionary simulations to investigate how and why the success of various strategies changes with the level of transparency. We demonstrate that non-zero transparency greatly reduces the set of successful memory-one strategies compared to the simultaneous iPD. For low and moderate transparency the classical “Win – Stay, Lose – Shift” (WSLS) strategy is the only evolutionary successful strategy. For high transparency all strategies are evolutionary unstable in the sense that they can be easily counteracted, and, finally, for maximal transparency a novel “Leader-Follower” strategy outperforms WSLS. Our results provide a partial explanation for the fact that the strategies proposed for the simultaneous iPD are rarely observed in nature, where high levels of transparency are common.

For humans and for non-human primates heart rate is a reliable indicator of an individual's current physiological state, with applications ranging from health checks to experimental studies of cognitive and emotional state. In humans, changes in the optical properties of the skin tissue correlated with cardiac cycles (imaging photoplethysmogram, iPPG) allow non-contact estimation of heart rate by its proxy, pulse rate. Yet, there is no established simple and non-invasive technique for pulse rate measurements in awake and behaving animals. Using iPPG, we here demonstrate that pulse rate in rhesus monkeys can be accurately estimated from facial videos. We computed iPPGs from eight color facial videos of four awake head-stabilized rhesus monkeys. Pulse rate estimated from iPPGs was in good agreement with reference data from a contact pulse-oximeter: the error of pulse rate estimation was below 5% of the individual average pulse rate in 83% of the epochs; the error was below 10% for 98% of the epochs. We conclude that iPPG allows non-invasive and non-contact estimation of pulse rate in non-human primates, which is useful for physiological studies and can be used toward welfare-assessment of non-human primates in research.

Highlights • Parkinson patients with hallucinations exhibit more perceptual errors. • Perceptual errors are due to decrease in perceptual sensitivity and not to response bias. • Parkinson patients with hallucinations exhibit increased performance fluctuations. • Deficits correlate with functional connectivity changes in fronto-parietal networks., Patients with Parkinson's disease (PD) frequently suffer from visual misperceptions and hallucinations, which are difficult to objectify and quantify. We aimed to develop an image recognition task to objectify misperceptions and to assess performance fluctuations in PD patients with and without self-reported hallucinations. Thirty-two non-demented patients with Parkinson's disease (16 with and 16 without self-reported visual hallucinations) and 25 age-matched healthy controls (HC) were tested. Participants performed a dynamic image recognition task with real and scrambled images. We assessed misperception scores and intra-individual variability in recognition times. To gain insight into possible neural mechanisms related to misperceptions and performance fluctuations we correlated resting state network connectivity to the behavioral outcomes in a subsample of Parkinson's disease patients (N = 16). We found that PD patients with self-reported hallucinations (PD-VH) exhibited higher perceptual error rates, due to decreased perceptual sensitivity and not due to changed decision criteria. In addition, PD-VH patients exhibited higher intra-individual variability in recognition times than HC or PD-nonVH patients. Both, misperceptions and intra-individual variability were negatively correlated with resting state functional connectivity involving frontal and parietal brain regions, albeit in partly different subregions. Consistent with previous research suggesting that hallucinations arise from dysfunction in attentional networks, misperception scores correlated with reduced functional connectivity between the dorsal attention and salience network. Intra-individual variability correlated with decreased connectivity between somatomotor and right fronto-parietal networks. We conclude that our task can detect visual misperceptions that are more prevalent in PD-VH patients. In addition, fluctuating visual performance appear to be a signature of PD-VH patients, which might assist further studies of the underlying pathophysiological mechanisms and cognitive processes.

Real-world agents, such as humans, animals and robots, observe each other during interactions and choose their own actions taking the partners’ ongoing behaviour into account. Yet, classical game theory assumes that players act either strictly sequentially or strictly simultaneously (without knowing the choices of each other). To account for action visibility and provide a more realistic model of interactions under time constraints, we introduce a new game-theoretic setting called transparent game, where each player has a certain probability to observe the choice of the partner before deciding on its own action. Using evolutionary simulations, we demonstrate that even a small probability of seeing the partner’s choice before one’s own decision substantially changes evolutionary successful strategies. Action visibility enhances cooperation in a Bach-or-Stravinsky game, but disrupts cooperation in a more competitive iterated Prisoner’s Dilemma. In both games, strategies based on the “Win–stay, lose–shift” and “Tit-for-tat” principles are predominant for moderate transparency, while for high transparency strategies of “Leader-Follower” type emerge. Our results have implications for studies of human and animal social behaviour, especially for the analysis of dyadic and group interactions.

Effort constitutes a major part of cost-benefit calculations underlying decision making. Therefore, estimating the effort someone has spent on a task is a core dimension for evaluating own and others’ actions. It has been previously shown that self-judgments of effort are influenced by the magnitude of obtained rewards. It is unclear, however, whether the influence of reward on effort estimations is limited to self-judgments or whether reward incorporation represents a general computational principle when judging effort. Here we show that people also integrate reward magnitude when judging the effort exerted by others. Participants (N=48) performed an effortful sensorimotor task interleaved with a partner, while rating either their own or the other person’s effort. After each trial but before the effort rating, both participants were informed about the obtained reward. We found that higher rewards led to higher estimations of exerted effort, in self-as well as other-judgments, and this effect was more pronounced for other-judgments. In both types of judgment, computational modelling revealed that reward information and the perceived level of exertion were combined in a Bayes optimal manner to form effort estimates. Remarkably, the extent to which rewards influenced effort judgments was positively correlated with conservative world-views, indicating that the basic computations underlying this behavioural phenomenon might be related to more general beliefs about the association between effort and reward in the society. The integration of reward information into retrospective effort judgments underscores the convergence of multiple information sources that supports adaptive learning and decision making in social contexts.

For humans and for non-human primates heart rate is a reliable indicator of an individual's current physiological state, with applications ranging from health checks to experimental studies of cognitive and emotional state. In humans, changes in the optical properties of the skin tissue correlated with cardiac cycles (imaging photoplethysmogram, iPPG) allow non-contact estimation of heart rate by its proxy, pulse rate. Yet, there is no established simple and non-invasive technique for pulse rate measurements in awake and behaving animals. Using iPPG, we here demonstrate that pulse rate in rhesus monkeys can be accurately estimated from facial videos. We computed iPPGs from eight color facial videos of four awake head-stabilized rhesus monkeys. Pulse rate estimated from iPPGs was in good agreement with reference data from a contact pulse-oximeter: the error of pulse rate estimation was below 5% of the individual average pulse rate in 83% of the epochs; the error was below 10% for 98% of the epochs. We conclude that iPPG allows non-invasive and non-contact estimation of pulse rate in non-human primates, which is useful for physiological studies and can be used toward welfare-assessment of non-human primates in research.

The present study aimed at investigating whether associated motivational salience causes preferential processing of inherently neutral faces similar to emotional expressions by means of event-related brain potentials (ERPs) and changes of the pupil size. To this aim, neutral faces were implicitly associated with monetary outcome, while participants (N = 44) performed a face-matching task with masked primes that ensured performance around chance level and thus an equal proportion of gain, loss, and zero outcomes. During learning, motivational context strongly impacted the processing of the fixation, prime and mask stimuli prior to the target face, indicated by enhanced amplitudes of subsequent ERP components and increased pupil size. In a separate test session, previously associated faces as well as novel faces with emotional expressions were presented within the same task but without motivational context and performance feedback. Most importantly, previously gain-associated faces amplified the LPC, although the individually contingent face-outcome assignments were not made explicit during the learning session. Emotional expressions impacted the N170 and EPN components. Modulations of the pupil size were absent in both motivationally-associated and emotional conditions. Our findings demonstrate that neural representations of neutral stimuli can acquire increased salience via implicit learning, with an advantage for gain over loss associations.

Humans and other animals constantly evaluate their decisions in order to learn and behave adaptively. Experimentally, such evaluation processes are accessed using metacognitive reports made after decisions, typically using verbally formulated confidence scales. When subjects report high confidence, it reflects a high certainty of being correct, but a low confidence might signify either low certainty about the outcome, or a high certainty of being incorrect. Hence, metacognitive reports might reflect not only different levels of decision certainty, but also two certainty directions (certainty of being correct and certainty of being incorrect). It is important to test if such bi-directional processing can be measured because, for decision-making under uncertainty, information about being incorrect is as important as information about being correct for guidance of subsequent behavior. We were able to capture implicit bi-directional certainty readouts by asking subjects to bet money on their perceptual decision accuracy using a six-grade wager scale (post-decision wagering, PDW). To isolate trial-specific aspects of metacognitive judgments, we used pre-decision wagering (wagering before the perceptual decision) to subtract, from PDW trials, influences resulting from non-trial-specific assessment of expected difficulty and psychological biases. This novel design allowed independent quantification of certainty of being correct and certainty of being incorrect, showing that subjects were able to read out certainty in a bi-directional manner. Certainty readouts about being incorrect were particularly associated with metacognitive sensitivity exceeding perceptual sensitivity (i.e. meta-d′ > d′), suggesting that such enhanced metacognitive efficiency is driven by information about incorrect decisions. Readouts of certainty in both directions increased on easier trials, and both certainty directions were also associated with faster metacognitive reaction times, indicating that certainty of being incorrect was not confounded with low certainty. Finally, both readouts influenced the amount of money subjects earned through PDW, suggesting that bi-directional readouts are important for planning future actions when feedback about previous decisions is unavailable.

Contralateral hemispheric representation of sensory inputs (the right visual hemifield in the left hemisphere and vice versa) is a fundamental feature of primate sensorimotor organization, in particular the visuomotor system. However, many higher-order cognitive functions in humans show an asymmetric hemispheric lateralization—e.g., right brain specialization for spatial processing—necessitating a convergence of information from both hemifields. Electrophysiological studies in monkeys and functional imaging in humans have investigated space and action representations at different stages of visuospatial processing, but the transition from contralateral to unified global spatial encoding and the relationship between these encoding schemes and functional lateralization are not fully understood. Moreover, the integration of data across monkeys and humans and elucidation of interspecies homologies is hindered, because divergent findings may reflect actual species differences or arise from discrepancies in techniques and measured signals (electrophysiology vs. imaging). Here, we directly compared spatial cue and memory representations for action planning in monkeys and humans using event-related functional MRI during a working-memory oculomotor task. In monkeys, cue and memory-delay period activity in the frontal, parietal, and temporal regions was strongly contralateral. In putative human functional homologs, the contralaterality was significantly weaker, and the asymmetry between the hemispheres was stronger. These results suggest an inverse relationship between contralaterality and lateralization and elucidate similarities and differences in human and macaque cortical circuits subserving spatial awareness and oculomotor goal-directed actions.

The posterior parietal cortex (PPC) has traditionally been considered important for awareness, spatial perception, and attention. However, recent findings provide evidence that the PPC also encodes information important for making decisions. These findings have initiated a running argument of whether the PPC is critically involved in decision making. To examine this issue, we reversibly inactivated the parietal reach region (PRR), the area of the PPC that is specialized for reaching movements, while two monkeys performed a memory-guided reaching or saccade task. The task included choices between two equally rewarded targets presented simultaneously in opposite visual fields. Free-choice trials were interleaved with instructed trials, in which a single cue presented in the peripheral visual field defined the reach and saccade target unequivocally. We found that PRR inactivation led to a strong reduction of contralesional choices, but only for reaches. On the other hand, saccade choices were not affected by PRR inactivation. Importantly, reaching and saccade movements to single instructed targets remained largely intact. These results cannot be explained as an effector-nonspecific deficit in spatial attention or awareness, since the temporary "lesion" had an impact only on reach choices. Hence, the PPR is a part of a network for reach decisions and not just reach planning.There has been an ongoing debate on whether the posterior parietal cortex (PPC) represents only spatial awareness, perception, and attention or whether it is also involved in decision making for actions. In this study we explore whether the parietal reach region (PRR), the region of the PPC that is specialized for reaches, is involved in the decision process. We inactivated the PRR while two monkeys performed reach and saccade choices between two targets presented simultaneously in both hemifields. We found that inactivation affected only the reach choices, while leaving saccade choices intact. These results cannot be explained as a deficit in attention, since the temporary lesion affected only the reach choices. Thus, PRR is a part of a network for making reach decisions.

Manipulation of the trunk midline has been shown to improve visuospatial performance in patients with unilateral visual neglect. The goal of the present study was to disentangle motor and perceptual components of egocentric midline manipulations and to investigate the contribution of individual hand preference. Two versions of visual temporal order judgment (TOJ) tasks were tested in healthy right- and left-handed subjects while trunk rotation was varied. In the congruent version, subjects were required to execute a saccade to the first of two horizontal stimuli presented with different stimulus onset asynchronies (SOA). In the incongruent version, subjects were required to perform a vertical saccade to a pre-learned color target, thereby dissociating motor response from the perceptual stimulus location. The main findings of this study are a trunk rotation and response direction specific impact on temporal judgments in form of a prior entry bias for right hemifield stimuli during rightward trunk rotation, but only in the congruent task. This trunk rotation-induced spatial bias was most pronounced in left-handed participants but had the same sign in the right-handed group. Results suggest that egocentric midline shifts in healthy subjects induce a spatially-specific motor, but not a perceptual, bias and underline the importance of taking individual differences in functional laterality such as handedness and mode of perceptual report into account when evaluating effects of trunk rotation in either healthy subjects or neurological patients.

Humans and other species continually perform microscopic eye movements, even when attending to a single point [1-3]. These movements, which include microscopic drifts and microsaccades, are under control of the oculomotor system [2, 4, 5], elicit strong responses throughout the visual system [6-11], and have been thought to serve important functions [12-16]. The influence of these fixational eye movements on the acquisition and neural processing of visual information remains unknown. Here, we show that during viewing of natural scenes, microscopic eye movements carry out a crucial information-processing step: they remove predictable correlations in natural scenes by equalizing the spatial power of the retinal image within the frequency range of ganglion cells' peak sensitivity. This transformation, which had been attributed to center-surround receptive field organization [17-19], occurs prior to any neural processing, and reveals a form of matching between the statistics of natural images and those of normal eye movements. We further show that the combined effect of microscopic eye movements and retinal receptive field organization is to convert spatial luminance discontinuities into synchronous firing events, thus beginning the process of edge extraction. In sum, our results show that microscopic eye movements are fundamental to two goals of early visual processing —redundancy reduction [20, 21] and feature extraction— and, thus, that neural representations are intrinsically sensory-motor from the very first processing stages.

Although neuronal responses in behaving monkeys are typically studied while the monkey fixates straight ahead, it is known that eye position modulates responses of visual neurons. The modulation has been found to enhance neuronal responses when the receptive field is placed in the straight-ahead position for neurons receiving input from the peripheral but not the central retina. We studied the effect of eye position on the responses of V1 complex cells receiving input from the central retina (1.1-5.7° eccentricity) while minimizing the effect of fixational eye movements. Contrast response functions were obtained separately with drifting light and dark bars. Data were fit with the Naka-Rushton equation: r(c)=Rmax×c/(c+c50)+s, where r(c) is mean spike rate at contrast c, Rmax is the maximum response, c50 is the contrast that elicits half of Rmax, and s is the spontaneous activity. Contrast sensitivity as measured by c50 was not affected by eye position. For dark bars, there was a statistically significant decline in the normalized Rmax with increasing deviation from straight ahead. Data for bright bars showed a similar trend with a less rapid decline. Our results indicate that neurons representing the central retina show a bias for the straight-ahead position resulting from modulation of the response gain without an accompanying modulation of contrast sensitivity. The modulation is especially obvious for dark stimuli, which might be useful for directing attention to hazardous situations such as dark holes or shadows concealing important objects (Supplement 1: Video Abstract, Supplemental digital content 1, http://links.lww.com/WNR/A295).

Humans rely on the fovea, the small region of the retina where receptors are most densely packed, for seeing fine spatial detail. Outside the fovea, it is well established that a variety of visual functions progressively decline with eccentricity [1–5]. In contrast, little is known about how vision varies within the central fovea, as incessant microscopic eye movements prevent isolation of adjacent foveal locations [6–8]. Using a new method for restricting visual stimulation to a selected retinal region, we examined the discrimination of fine patterns at different eccentricities within the foveola. We show that high-acuity judgments are impaired when stimuli are presented just a few arcminutes away from the preferred retinal locus of fixation. Furthermore, we show that this dependence on eccentricity is normally counter-balanced by the occurrence of precisely directed microsaccades, which bring the preferred fixation locus onto the stimulus. Thus, contrary to common assumptions, vision is not uniform within the foveola, but targeted microscopic eye movements compensate for this lack of homogeneity. Our results reveal that microsaccades, like larger saccades, enable examination of the stimulus at a finer level of detail and suggest that a reduced precision in oculomotor control may be responsible for the visual acuity impairments observed in various disorders.

The ability to selectively process visual inputs and to decide between multiple movement options in an adaptive manner is critical for survival. Such decisions are known to be influenced by factors such as reward expectation and visual saliency. The dorsal pulvinar connects to a multitude of cortical areas that are involved in visuospatial memory and integrate information about upcoming eye movements with expected reward values. However, it is unclear whether the dorsal pulvinar is critically involved in spatial memory and reward-based oculomotor decision behavior. To examine this, we reversibly inactivated the dorsal portion of the pulvinar while monkeys performed a delayed memory saccade task that included choices between equally or unequally rewarded options. Pulvinar inactivation resulted in a delay of saccade initiation toward memorized contralesional targets but did not affect spatial memory. Furthermore, pulvinar inactivation caused a pronounced choice bias toward the ipsilesional hemifield when the reward value in the two hemifields was equal. However, this choice bias could be alleviated by placing a high reward target into the contralesional hemifield. The bias was less affected by the manipulation of relative visual saliency between the two competing targets. These results suggest that the dorsal pulvinar is involved in determining the behavioral desirability of movement goals while being less critical for spatial memory and reward processing.