Dorsal pulvinar inactivation leads to spatial selection bias without perceptual deficit.

The dorsal pulvinar has been implicated in visuospatial attentional and perceptual confidence processing. Pulvinar lesions in humans and monkeys lead to spatial neglect symptoms, including an overt spatial saccade bias during free choices. However, it remains unclear whether disrupting the dorsal pulvinar during target selection that relies on a perceptual decision leads to a perceptual impairment or a more general spatial orienting and choice deficit. To address this question, we reversibly inactivated the unilateral dorsal pulvinar by injecting GABA-A agonist THIP while two macaque monkeys performed a color discrimination saccade task with varying perceptual difficulty. We used Signal Detection Theory and simulations to dissociate perceptual sensitivity (d-prime) and spatial selection bias (response criterion) effects. We expected a decrease in d-prime if dorsal pulvinar affects perceptual discrimination and a shift in response criterion if dorsal pulvinar is mainly involved in spatial orienting. After the inactivation, we observed response criterion shifts away from contralesional stimuli, especially when two competing stimuli in opposite hemifields were present. Notably, the d-prime and overall accuracy remained largely unaffected. Our results underline the critical contribution of the dorsal pulvinar to spatial orienting and action selection while showing it to be less important for visual perceptual discrimination.

Visual, delay, and oculomotor timing and tuning in macaque dorsal pulvinar during instructed and free choice memory saccades.

Causal perturbations suggest that primate dorsal pulvinar plays a crucial role in target selection and saccade planning, though its basic neuronal properties remain unclear. Some functional aspects of dorsal pulvinar and interconnected frontoparietal areas-e.g. ipsilesional choice bias after inactivation-are similar. But it is unknown if dorsal pulvinar shares oculomotor properties of cortical circuitry, in particular delay and choice-related activity. We investigated such properties in macaque dorsal pulvinar during instructed and free-choice memory saccades. Most recorded units showed visual (12%), saccade-related (30%), or both types of responses (22%). Visual responses were primarily contralateral; diverse saccade-related responses were predominantly post-saccadic with a weak contralateral bias. Memory delay and pre-saccadic enhancement was infrequent (11-9%)-instead, activity was often suppressed during saccade planning (25%) and further during execution (15%). Surprisingly, only few units exhibited classical visuomotor patterns combining cue and continuous delay activity or pre-saccadic ramping; moreover, 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 visual and post-saccadic responses imply its participation in integrating spatial goals with processing across saccades.

Detection of Transcranial Alternating Current Stimulation Aftereffects Is Improved by Considering the Individual Electric Field Strength and Self-Rated Sleepiness.

Non-invasive electrical stimulation methods, such as transcranial alternating current stimulation (tACS), are increasingly used in human neuroscience research and offer potential new avenues to treat neurological and psychiatric disorders. However, their often variable effects have also raised concerns in the scientific and clinical communities. This study aims to investigate the influence of subject-specific factors on the alpha tACS-induced aftereffect on the alpha amplitude (measured with electroencephalography, EEG) as well as on the connectivity strength between nodes of the default mode network (DMN) [measured with functional magnetic resonance imaging (fMRI)]. As subject-specific factors we considered the individual electrical field (EFIELD) strength at target regions in the brain, the frequency mismatch between applied stimulation and individual alpha frequency (IAF) and as a covariate, subject's changes in mental state, i.e., sleepiness. Eighteen subjects participated in a tACS and a sham session conducted on different days. Each session consisted of three runs (pre/stimulation/). tACS was applied during the second run at each subject's individual alpha frequency (IAF), applying 1 mA peak-to-peak intensity for 7 min, using an occipital bihemispheric montage. In every run, subjects watched a video designed to increase in-scanner compliance. To investigate the aftereffect of tACS on EEG alpha amplitude and on DMN connectivity strength, EEG data were recorded simultaneously with fMRI data. Self-rated sleepiness was documented using a questionnaire. Conventional statistics (ANOVA) did not show a significant aftereffect of tACS on the alpha amplitude compared to sham stimulation. Including individual EFIELD strengths and self-rated sleepiness scores in a multiple linear regression model, significant tACS-induced aftereffects were observed. However, the subject-wise mismatch between tACS frequency and IAF had no contribution to our model. Neither standard nor extended statistical methods confirmed a tACS-induced aftereffect on DMN functional connectivity. Our results show that it is possible and necessary to disentangle alpha amplitude changes due to intrinsic mechanisms and to external manipulation using tACS on the alpha amplitude that might otherwise be overlooked. Our results suggest that EFIELD is really the most significant factor that explains the alpha amplitude modulation during a tACS session. This knowledge helps to understand the variability of the tACS-induced aftereffects.

The effect of subliminal incentives on goal-directed eye movements.

Over the last decades, several studies have demonstrated that conscious and unconscious reward incentives both affect performance in physical and cognitive tasks, suggesting that goal pursuit can arise from an unconscious will. Whether the planning of goal-directed saccadic eye movements during an effortful task can also be affected by subliminal reward cues has not been systematically investigated. We employed a novel task where participants made several eye movements back and forth between a fixation point and a number of peripheral targets. The total number of targets visited by the eyes in a fixed amount of time determined participants' monetary gain. The magnitude of the reward at stake was briefly shown at the beginning of each trial and masked by pattern images superimposed in time so that at shorter display durations participants perceived reward incentives subliminally. We found a main effect of reward across all display durations as higher reward enhanced participants' oculomotor effort measured as the frequency and peak velocity of saccades. This effect was strongest for consciously perceived rewards but also occurred when rewards were subliminally perceived. Although we did not find a statistically significant dissociation between the reward-related modulation of different saccadic parameters, across two experiments the most robust effect of subliminal rewards was observed for the modulation of the saccadic frequency but not the peak velocity. These results suggest that multiple indices of oculomotor effort can be incentivized by subliminal rewards and that saccadic frequency may provide the most sensitive indicator of subliminal incentivization of eye movements. NEW & NOTEWORTHY Reward incentives motivate humans to exert more effort, and they do so even when rewards are subconsciously perceived. It has been unknown whether these effects also extend to eye movements that have lower energetic demands compared with other movement types. We devised a behavioral task that required fast execution of multiple eye movements. Subliminal rewards enhanced the frequency and peak velocity of saccadic eye movements, with the most reliable effect observed for saccadic frequency.

Effective connectivity and spatial selectivity-dependent fMRI changes elicited by microstimulation of pulvinar and LIP.

The thalamic pulvinar and the lateral intraparietal area (LIP) share reciprocal anatomical connections and are part of an extensive cortical and subcortical network involved in spatial attention and oculomotor processing. The goal of this study was to compare the effective connectivity of dorsal pulvinar (dPul) and LIP and to probe the dependency of microstimulation effects on task demands and spatial tuning properties of a given brain region. To this end, we applied unilateral electrical microstimulation in the dPul (mainly medial pulvinar) and LIP in combination with event-related BOLD fMRI in monkeys performing fixation and memory-guided saccade tasks. Microstimulation in both dPul and LIP enhanced task-related activity in monosynaptically-connected fronto-parietal cortex and along the superior temporal sulcus (STS) including putative face patch locations, as well as in extrastriate cortex. LIP microstimulation elicited strong activity in the opposite homotopic LIP while no homotopic activation was found with dPul stimulation. Both dPul and LIP stimulation also elicited activity in several heterotopic cortical areas in the opposite hemisphere, implying polysynaptic propagation of excitation. Despite extensive activation along the intraparietal sulcus evoked by LIP stimulation, there was a difference in frontal and occipital connectivity elicited by posterior and anterior LIP stimulation sites. Comparison of dPul stimulation with the adjacent but functionally dissimilar ventral pulvinar also showed distinct connectivity. On the level of single trial timecourses within each region of interest (ROI), most ROIs did not show task-dependence of stimulation-elicited response modulation. Across ROIs, however, there was an interaction between task and stimulation, and task-specific correlations between the initial spatial selectivity and the magnitude of stimulation effect were observed. Consequently, stimulation-elicited modulation of task-related activity was best fitted by an additive model scaled down by the initial response amplitude. In summary, we identified overlapping and distinct patterns of thalamocortical and corticocortical connectivity of pulvinar and LIP, highlighting the dorsal bank and fundus of STS as a prominent node of shared circuitry. Spatial task-specific and partly polysynaptic modulations of cue and saccade planning delay period activity in both hemispheres exerted by unilateral pulvinar and parietal stimulation provide insight into the distributed interhemispheric processing underlying spatial behavior.

Reduced alpha amplitudes predict perceptual suppression.

The amplitude of prestimulus alpha oscillations over parieto-occipital cortex has been shown to predict visual detection of masked and threshold-level stimuli. Whether alpha activity similarly predicts target visibility in perceptual suppression paradigms, another type of illusion commonly used to investigate visual awareness, is presently unclear. Here, we examined prestimulus alpha activity in the electroencephalogram (EEG) of healthy participants in the context of a generalized flash suppression (GFS) task during which salient target stimuli are rendered subjectively invisible in a subset of trials following the onset of a full-field motion stimulus. Unlike for masking or threshold paradigms, alpha (8-12 Hz) amplitude prior to motion onset was significantly higher when targets remained subjectively visible compared to trials during which the targets became perceptually suppressed. Furthermore, individual prestimulus alpha amplitudes strongly correlated with the individual trial-to-trial variability quenching following motion stimulus onset, indicating that variability quenching in visual cortex is closely linked to prestimulus alpha activity. We conclude that predictive correlates of conscious perception derived from perceptual suppression paradigms differ substantially from those obtained with "near threshold paradigms", possibly reflecting the effectiveness of the suppressor stimulus.

Combining brain perturbation and neuroimaging in non-human primates.

Brain perturbation studies allow detailed causal inferences of behavioral and neural processes. Because the combination of brain perturbation methods and neural measurement techniques is inherently challenging, research in humans has predominantly focused on non-invasive, indirect brain perturbations, or neurological lesion studies. Non-human primates have been indispensable as a neurobiological system that is highly similar to humans while simultaneously being more experimentally tractable, allowing visualization of the functional and structural impact of systematic brain perturbation. This review considers the state of the art in non-human primate brain perturbation with a focus on approaches that can be combined with neuroimaging. We consider both non-reversible (lesions) and reversible or temporary perturbations such as electrical, pharmacological, optical, optogenetic, chemogenetic, pathway-selective, and ultrasound based interference methods. Method-specific considerations from the research and development community are offered to facilitate research in this field and support further innovations. We conclude by identifying novel avenues for further research and innovation and by highlighting the clinical translational potential of the methods.

Author Correction: Thalamus exhibits less sensory variability quenching than cortex.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Extracting Robust Biomarkers From Multichannel EEG Time Series Using Nonlinear Dimensionality Reduction Applied to Ordinal Pattern Statistics and Spectral Quantities.

In this study, ordinal pattern analysis and classical frequency-based EEG analysis methods are used to differentiate between EEGs of different age groups as well as individuals. As characteristic features, functional connectivity as well as single-channel measures in both the time and frequency domain are considered. We compare the separation power of each feature set after nonlinear dimensionality reduction using t-distributed stochastic neighbor embedding and demonstrate that ordinal pattern-based measures yield results comparable to frequency-based measures applied to preprocessed data, and outperform them if applied to raw data. Our analysis yields no significant differences in performance between single-channel features and functional connectivity features regarding the question of age group separation.

Probing the Link Between Perception and Oscillations: Lessons from Transcranial Alternating Current Stimulation.

Brain oscillations are regarded as important for perception as they open and close time windows for neural spiking to enable the effective communication within and across brain regions. In the past, studies on perception primarily relied on the use of electrophysiological techniques for probing a correlative link between brain oscillations and perception. The emergence of noninvasive brain stimulation techniques such as transcranial alternating current stimulation (tACS) provides the possibility to study the causal contribution of specific oscillatory frequencies to perception. Here, we review the studies on visual, auditory, and somatosensory perception that employed tACS to probe the causality of brain oscillations for perception. The current literature is consistent with a causal role of alpha and gamma oscillations in parieto-occipital regions for visual perception and theta and gamma oscillations in auditory cortices for auditory perception. In addition, the sensory gating by alpha oscillations applies not only to the visual but also to the somatosensory domain. We conclude that albeit more refined perceptual paradigms and individualized stimulation practices remain to be systematically adopted, tACS is a promising tool for establishing a causal link between neural oscillations and perception.

Aberrant functional connectivity of resting state networks related to misperceptions and intra-individual variability in Parkinson's disease.

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.

Eye position signals in the dorsal pulvinar during fixation and goal-directed saccades.

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.

Trunk rotation and handedness modulate cortical activation in neglect-associated regions during temporal order judgments.

The rotation of the trunk around its vertical midline could be shown to bias visuospatial temporal judgments towards targets in the hemifield ipsilateral to the trunk orientation and to improve visuospatial performance in patients with visual neglect. However, the underlying brain mechanisms are not well understood. Therefore, the goal of the present study was to investigate the neural effects associated with egocentric midplane shifts under consideration of individual handedness. We employed a visuospatial temporal order judgment (TOJ) task in healthy right- and left-handed subjects while their trunk rotation was varied. Participants responded by a saccade towards the stimulus perceived first out of two stimuli presented with different stimulus onset asynchronies (SOA). Apart from gaze behavior, BOLD-fMRI responses were measured using functional magnetic resonance imaging (fMRI). Based on findings from spatial neglect research, analyses of fMRI-BOLD responses were focused on a bilateral fronto-temporo-parietal network comprising Brodmann areas 22, 39, 40, and 44, as well as the basal ganglia core nuclei (caudate, putamen, pallidum). We observed an acceleration of saccadic speed towards stimuli ipsilateral to the trunk orientation modulated by individual handedness. Left-handed participants showed the strongest behavioral and neural effects, suggesting greater susceptibility to manipulations of trunk orientation. With respect to the dominant hand, a rotation around the vertical trunk midline modulated the activation of an ipsilateral network comprising fronto-temporo-parietal regions and the putamen with the strongest effects for saccades towards the hemifield opposite to the dominant hand. Within the investigated network, the temporo-parietal junction (TPJ) appears to serve as a region integrating sensory, motor, and trunk position information. Our results are discussed in the context of gain modulatory and laterality effects.

Thalamus exhibits less sensory variability quenching than cortex.

Spiking activity exhibits a large degree of variability across identical trials, which has been shown to be significantly reduced by stimulus onset in a wide range of cortical areas. Whether similar dynamics apply to the thalamus and in particular to the pulvinar is largely unknown. Here, we examined electrophysiological recordings from two adult rhesus macaques performing a perceptual task and comparatively investigated trial-to-trial variability in higher-order thalamus (ventral and dorsal pulvinar), the lateral geniculate nucleus (LGN) and visual cortex (area V4) prior to and following the presentation of a visual stimulus. We found spiking variability during stable fixation prior to stimulus onset to be considerably lower in both pulvinar and the LGN as compared to area V4. In contrast to the prominent variability reduction in V4 upon stimulus onset, variability in the thalamic nuclei was largely unaffected by visual stimulation. There was a small but significant variability decrease in the dorsal pulvinar, but not in the ventral portion of the pulvinar, which is closely connected to visual cortices and would thus have been expected to reflect cortical response properties. This dissociation did not stem from differences in response strength or mean firing rates and indicates fundamental differences in variability quenching between thalamus and cortex.

Post-decision wagering after perceptual judgments reveals bi-directional certainty readouts.

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.

Reach and grasp deficits following damage to the dorsal pulvinar.

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.

Consciousness Regained: Disentangling Mechanisms, Brain Systems, and Behavioral Responses.

How consciousness (experience) arises from and relates to material brain processes (the "mind-body problem") has been pondered by thinkers for centuries, and is regarded as among the deepest unsolved problems in science, with wide-ranging theoretical, clinical, and ethical implications. Until the last few decades, this was largely seen as a philosophical topic, but not widely accepted in mainstream neuroscience. Since the 1980s, however, novel methods and theoretical advances have yielded remarkable results, opening up the field for scientific and clinical progress. Since a seminal paper by Crick and Koch (1998) claimed that a science of consciousness should first search for its neural correlates (NCC), a variety of correlates have been suggested, including both content-specific NCCs, determining particular phenomenal components within an experience, and the full NCC, the neural substrates supporting entire conscious experiences. In this review, we present recent progress on theoretical, experimental, and clinical issues. Specifically, we (1) review methodological advances that are important for dissociating conscious experience from related enabling and executive functions, (2) suggest how critically reconsidering the role of the frontal cortex may further delineate NCCs, (3) advocate the need for general, objective, brain-based measures of the capacity for consciousness that are independent of sensory processing and executive functions, and (4) show how animal studies can reveal population and network phenomena of relevance for understanding mechanisms of consciousness.

Simultaneous Transcranial Alternating Current Stimulation and Functional Magnetic Resonance Imaging.

Transcranial alternating current stimulation (tACS) is a promising tool for noninvasive investigation of brain oscillations. TACS employs frequency-specific stimulation of the human brain through current applied to the scalp with surface electrodes. Most current knowledge of the technique is based on behavioral studies; thus, combining the method with brain imaging holds potential to better understand the mechanisms of tACS. Because of electrical and susceptibility artifacts, combining tACS with brain imaging can be challenging, however, one brain imaging technique that is well suited to be applied simultaneously with tACS is functional magnetic resonance imaging (fMRI). In our lab, we have successfully combined tACS with simultaneous fMRI measurements to show that tACS effects are state, current, and frequency dependent, and that modulation of brain activity is not limited to the area directly below the electrodes. This article describes a safe and reliable setup for applying tACS simultaneously with visual task fMRI studies, which can lend to understanding oscillatory brain function as well as the effects of tACS on the brain.

Electrical Microstimulation of the Pulvinar Biases Saccade Choices and Reaction Times in a Time-Dependent Manner.

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 STATEMENT Despite 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.