Dorsal attention network activity during perceptual organization is distinct in schizophrenia and predictive of cognitive disorganization.

Visual shape completion is a canonical perceptual organization process that integrates spatially distributed edge information into unified representations of objects. People with schizophrenia show difficulty in discriminating completed shapes, but the brain networks and functional connections underlying this perceptual difference remain poorly understood. Also unclear is whether brain network differences in schizophrenia occur in related illnesses or vary with illness features transdiagnostically. To address these topics, we scanned (functional magnetic resonance imaging, fMRI) people with schizophrenia, bipolar disorder, or no psychiatric illness during rest and during a task in which they discriminated configurations that formed or failed to form completed shapes (illusory and fragmented condition, respectively). Multivariate pattern differences were identified on the cortical surface using 360 predefined parcels and 12 functional networks composed of such parcels. Brain activity flow mapping was used to evaluate the likely involvement of resting-state connections for shape completion. Illusory/fragmented task activation differences ('modulations') in the dorsal attention network (DAN) could distinguish people with schizophrenia from the other groups (AUCs > .85) and could transdiagnostically predict cognitive disorganization severity. Activity flow over functional connections from the DAN could predict secondary visual network modulations in each group, except in schizophrenia. The secondary visual network was strongly and similarly modulated in each group. Task modulations were dispersed over more networks in patients compared to controls. In summary, DAN activity during visual perceptual organization is distinct in schizophrenia, symptomatically relevant, and potentially related to improper attention-related feedback into secondary visual areas.

Brain network mechanisms of visual shape completion.

Visual shape completion recovers object shape, size, and number from spatially segregated edges. Despite being extensively investigated, the process's underlying brain regions, networks, and functional connections are still not well understood. To shed light on the topic, we scanned (fMRI) healthy adults during rest and during a task in which they discriminated pac-man configurations that formed or failed to form completed shapes (illusory and fragmented condition, respectively). Task activation differences (illusory-fragmented), resting-state functional connectivity, and multivariate patterns were identified on the cortical surface using 360 predefined parcels and 12 functional networks composed of such parcels. Brain activity flow mapping (ActFlow) was used to evaluate the likely involvement of resting-state connections for shape completion. We identified 36 differentially-active parcels including a posterior temporal region, PH, whose activity was consistent across 95% of observers. Significant task regions primarily occupied the secondary visual network but also incorporated the frontoparietal, dorsal attention, default mode, and cingulo-opercular networks. Each parcel's task activation difference could be modeled via its resting-state connections with the remaining parcels (r=.62, p<10-9), suggesting that such connections undergird shape completion. Functional connections from the dorsal attention network were key in modelling task activation differences in the secondary visual network. Dorsal attention and frontoparietal connections could also model activations in the remaining networks. Taken together, these results suggest that shape completion relies upon a sparsely distributed but densely interconnected network coalition that is centered in the secondary visual network, coordinated by the dorsal attention network, and inclusive of at least three other networks.

N-methyl d-aspartate receptor hypofunction reduces visual contextual integration.

Visual cognition is finely tuned to the elements in a scene but also relies on contextual integration to improve visual detection and discrimination. This integration is impaired in patients with schizophrenia. Studying impairments in contextual integration may lead to biomarkers of schizophrenia, tools to monitor disease progression, and, in animal models, insight into the underlying neural deficits. We developed a nonhuman primate model to test the hypothesis that hypofunction of the N-methyl d-aspartate receptor (NMDAR) impairs contextual integration. Two male rhesus macaques (Macaca mulatta) were trained to indicate which of two patterns on the screen had the highest contrast. One of these patterns appeared in isolation, and the other was surrounded by a high-contrast pattern. In humans, this high-contrast context is known to lead to an underestimation of contrast. This so-called Chubb illusion is thought to result from surround suppression, a key contextual integration mechanism. To test the involvement of NMDAR in this process, we compared animals' perceptual bias with and without intramuscular injections of a subanesthetic dose of the NMDAR antagonist ketamine. In the absence of ketamine, the animals reported a Chubb illusion - matching reports in healthy humans. Hence, monkeys - just like humans - perform visual contextual integration. This reaffirms the importance of nonhuman primates to help understand visual cognition. Injection of ketamine significantly reduced the strength of the illusion and thus impaired contextual integration. This supports the hypothesis that NMDAR hypofunction plays a causal role in specific behavioral impairments observed in schizophrenia.

Transcranial alternating current stimulation attenuates BOLD adaptation and increases functional connectivity.

Transcranial alternating current stimulation (tACS) is used as a noninvasive tool for cognitive enhancement and clinical applications. The physiological effects of tACS, however, are complex and poorly understood. Most studies of tACS focus on its ability to entrain brain oscillations, but our behavioral results in humans and extracellular recordings in nonhuman primates support the view that tACS at 10 Hz also affects brain function by reducing sensory adaptation. Our primary goal in the present study is to test this hypothesis using blood oxygen level-dependent (BOLD) imaging in human subjects. Using concurrent functional magnetic resonance imaging (fMRI) and tACS, and a motion adaptation paradigm developed to quantify BOLD adaptation, we show that tACS significantly attenuates adaptation in the human motion area (hMT+). In addition, an exploratory analysis shows that tACS increases functional connectivity of the stimulated hMT+ with the rest of the brain and the dorsal attention network in particular. Based on field estimates from individualized head models, we relate these changes to the strength of tACS-induced electric fields. Specifically, we report that functional connectivity (between hMT+ and any other region of interest) increases in proportion to the field strength in the region of interest. These findings add support for the claim that weak 10-Hz currents applied to the scalp modulate both local and global measures of brain activity.NEW & NOTEWORTHY Concurrent transcranial alternating current stimulation (tACS) and functional MRI show that tACS affects the human brain by attenuating adaptation and increasing functional connectivity in a dose-dependent manner. This work is important for our basic understanding of what tACS does, but also for therapeutic applications, which need insight into the full range of ways in which tACS affects the brain.

Transcranial Alternating Current Stimulation Attenuates Neuronal Adaptation.

We previously showed that brief application of 2 mA (peak-to-peak) transcranial currents alternating at 10 Hz significantly reduces motion adaptation in humans. This is but one of many behavioral studies showing that weak currents applied to the scalp modulate neural processing. Transcranial stimulation has been shown to improve perception, learning, and a range of clinical symptoms. Few studies, however, have measured the neural consequences of transcranial current stimulation. We capitalized on the strong link between motion perception and neural activity in the middle temporal (MT) area of the macaque monkey to study the neural mechanisms that underlie the behavioral consequences of transcranial alternating current stimulation. First, we observed that 2 mA currents generated substantial intracranial fields, which were much stronger in the stimulated hemisphere (0.12 V/m) than on the opposite side of the brain (0.03 V/m). Second, we found that brief application of transcranial alternating current stimulation at 10 Hz reduced spike-frequency adaptation of MT neurons and led to a broadband increase in the power spectrum of local field potentials. Together, these findings provide a direct demonstration that weak electric fields applied to the scalp significantly affect neural processing in the primate brain and that this includes a hitherto unknown mechanism that attenuates sensory adaptation.SIGNIFICANCE STATEMENT Transcranial stimulation has been claimed to improve perception, learning, and a range of clinical symptoms. Little is known, however, how transcranial current stimulation generates such effects, and the search for better stimulation protocols proceeds largely by trial and error. We investigated, for the first time, the neural consequences of stimulation in the monkey brain. We found that even brief application of alternating current stimulation reduced the effects of adaptation on single-neuron firing rates and local field potentials; this mechanistic insight explains previous behavioral findings and suggests a novel way to modulate neural information processing using transcranial currents. In addition, by developing an animal model to help understand transcranial stimulation, this study will aid the rational design of stimulation protocols for the treatment of mental illnesses, and the improvement of perception and learning.

Evidence and Counterevidence in Motion Perception.

Sensory neurons gather evidence in favor of the specific stimuli to which they are tuned, but they could improve their sensitivity by also taking counterevidence into account. The Bours-Lankheet model for motion detection uses counterevidence that relies on a specific combination of the ON and OFF channels in the early visual system. Specifically, the model detects pairs of flashes that occur separated in space and time. If the flashes have the same contrast polarity, they are interpreted as evidence in favor of the corresponding motion. But if they have opposite contrasts, they are interpreted as evidence against it. This mechanism provides an explanation for reverse-phi (the perceived reversal of an apparent motion stimulus due to periodic contrast-inversions) that is a conceptual departure from the standard explanations of the effect. Here, we investigate this counterevidence mechanism by measuring directional tuning curves of neurons in the primary visual and middle temporal cortex areas of awake, behaving macaques using constant-contrast and inverting-contrast moving dot stimuli. Our electrophysiological data support the Bours-Lankheet model and suggest that the counterevidence computation occurs at an early stage of neural processing not captured by the standard models.

Perisaccadic visual perception.

Primates use frequent, rapid eye movements to sample their visual environment. This is a fruitful strategy to make the best use of the highly sensitive foveal part of the retina, but it requires neural mechanisms to bind the rapidly changing visual input into a single, stable percept. Studies investigating these neural mechanisms have typically assumed that perisaccadic perception in nonhuman primates matches that of humans. We tested this assumption by performing identical experiments in human and nonhuman primates. Our data confirm that perisaccadic visual perception of macaques and humans is qualitatively similar. Specifically, we found a reduction in detectability and mislocalization of targets presented at the time of saccades. We also found substantial differences between human and nonhuman primates. Notably, in nonhuman primates, localization that requires knowledge of eye position was less precise, nonhuman primates detected fewer perisaccadic stimuli, and perisaccadic compression was not towards the saccade target. The qualitative similarities between species support the view that the nonhuman primate is ideally suited to study aspects of brain function-such as those relying on foveal vision-that are uniquely developed in primates. The quantitative differences, however, demonstrate the need for a reassessment of the models purportedly linking neural response changes at the time of saccades with the behavioral phenomena of perisaccadic reduction of detectability and mislocalization.

Transcranial alternating current stimulation attenuates visual motion adaptation.

Transcranial alternating current stimulation (tACS) is used in clinical applications and basic neuroscience research. Although its behavioral effects are evident from prior reports, current understanding of the mechanisms that underlie these effects is limited. We used motion perception, a percept with relatively well known properties and underlying neural mechanisms to investigate tACS mechanisms. Healthy human volunteers showed a surprising improvement in motion sensitivity when visual stimuli were paired with 10 Hz tACS. In addition, tACS reduced the motion-after effect, and this reduction was correlated with the improvement in motion sensitivity. Electrical stimulation had no consistent effect when applied before presenting a visual stimulus or during recovery from motion adaptation. Together, these findings suggest that perceptual effects of tACS result from an attenuation of adaptation. Important consequences for the practical use of tACS follow from our work. First, because this mechanism interferes only with adaptation, this suggests that tACS can be targeted at subsets of neurons (by adapting them), even when the applied currents spread widely throughout the brain. Second, by interfering with adaptation, this mechanism provides a means by which electrical stimulation can generate behavioral effects that outlast the stimulation.

Eye-position signals in the dorsal visual system are accurate and precise on short timescales.

Eye-position signals (EPS) are found throughout the primate visual system and are thought to provide a mechanism for representing spatial locations in a manner that is robust to changes in eye position. It remains unknown, however, whether cortical EPS (also known as "gain fields") have the necessary spatial and temporal characteristics to fulfill their purported computational roles. To quantify these EPS, we combined single-unit recordings in four dorsal visual areas of behaving rhesus macaques (lateral intraparietal area, ventral intraparietal area, middle temporal area, and the medial superior temporal area) with likelihood-based population-decoding techniques. The decoders used knowledge of spiking statistics to estimate eye position during fixation from a set of observed spike counts across neurons. Importantly, these samples were short in duration (100 ms) and from individual trials to mimic the real-time estimation problem faced by the brain. The results suggest that cortical EPS provide an accurate and precise representation of eye position, albeit with unequal signal fidelity across brain areas and a modest underestimation of eye eccentricity. The underestimation of eye eccentricity predicted a pattern of mislocalization that matches the errors made by human observers. In addition, we found that eccentric eye positions were associated with enhanced precision relative to the primary eye position. This predicts that positions in visual space should be represented more reliably during eccentric gaze than while looking straight ahead. Together, these results suggest that cortical eye-position signals provide a useable head-centered representation of visual space on timescales that are compatible with the duration of a typical ocular fixation.

Similar adaptation effects in primary visual cortex and area MT of the macaque monkey under matched stimulus conditions.

Recent stimulus history, or adaptation, can alter neuronal response properties. Adaptation effects have been characterized in a number of visually responsive structures, from the retina to higher visual cortex. However, it remains unclear whether adaptation effects across stages of the visual system take a similar form in response to a particular sensory event. This is because studies typically probe a single structure or cortical area, using a stimulus ensemble chosen to provide potent drive to the cells of interest. Here we adopt an alternative approach and compare adaptation effects in primary visual cortex (V1) and area MT using identical stimulus ensembles. Previous work has suggested these areas adjust to recent stimulus drive in distinct ways. We show that this is not the case: adaptation effects in V1 and MT can involve weak or strong loss of responsivity and shifts in neuronal preference toward or away from the adapter, depending on stimulus size and adaptation duration. For a particular stimulus size and adaptation duration, however, effects are similar in nature and magnitude in V1 and MT. We also show that adaptation effects in MT of awake animals depend strongly on stimulus size. Our results suggest that the strategies for adjusting to recent stimulus history depend more strongly on adaptation duration and stimulus size than on the cortical area. Moreover, they indicate that different levels of the visual system adapt similarly to recent sensory experience.

Transcranial direct current stimulation over posterior parietal cortex modulates visuospatial localization.

Visual localization is based on the complex interplay of bottom-up and top-down processing. Based on previous work, the posterior parietal cortex (PPC) is assumed to play an essential role in this interplay. In this study, we investigated the causal role of the PPC in visual localization. Specifically, our goal was to determine whether modulation of the PPC via transcranial direct current stimulation (tDCS) could induce visual mislocalization similar to that induced by an exogenous attentional cue (Wright, Morris, & Krekelberg, 2011). We placed one stimulation electrode over the right PPC and the other over the left PPC (dual tDCS) and varied the polarity of the stimulation. We found that this manipulation altered visual localization; this supports the causal involvement of the PPC in visual localization. Notably, mislocalization was more rightward when the cathode was placed over the right PPC than when the anode was placed over the right PPC. This mislocalization was found within a few minutes of stimulation onset, it dissipated during stimulation, but then resurfaced after stimulation offset and lasted for another 10-15 min. On the assumption that excitability is reduced beneath the cathode and increased beneath the anode, these findings support the view that each hemisphere biases processing to the contralateral hemifield and that the balance of activation between the hemispheres contributes to position perception (Kinsbourne, 1977; Szczepanski, Konen, & Kastner, 2010).

Neural mechanisms of speed perception: transparent motion.

Visual motion on the macaque retina is processed by direction- and speed-selective neurons in extrastriate middle temporal cortex (MT). There is strong evidence for a link between the activity of these neurons and direction perception. However, there is conflicting evidence for a link between speed selectivity of MT neurons and speed perception. Here we study this relationship by using a strong perceptual illusion in speed perception: when two transparently superimposed dot patterns move in opposite directions, their apparent speed is much larger than the perceived speed of a single pattern moving at that physical speed. Moreover, the sensitivity for speed discrimination is reduced for such bidirectional patterns. We first confirmed these behavioral findings in human subjects and extended them to a monkey subject. Second, we determined speed tuning curves of MT neurons to bidirectional motion and compared these to speed tuning curves for unidirectional motion. Consistent with previous reports, the response to bidirectional motion was often reduced compared with unidirectional motion at the preferred speed. In addition, we found that tuning curves for bidirectional motion were shifted to lower preferred speeds. As a consequence, bidirectional motion of some speeds typically evoked larger responses than unidirectional motion. Third, we showed that these changes in neural responses could explain changes in speed perception with a simple labeled line decoder. These data provide new insight into the encoding of transparent motion patterns and provide support for the hypothesis that MT activity can be decoded for speed perception with a labeled line model.

Intrasaccadic suppression is dominated by reduced detector gain.

Human vision requires fast eye movements (saccades). Each saccade causes a self-induced motion signal, but we are not aware of this potentially jarring visual input. Among the theorized causes of this phenomenon is a decrease in visual sensitivity before (presaccadic suppression) and during (intrasaccadic suppression) saccades. We investigated intrasaccadic suppression using a perceptual template model (PTM) relating visual detection to different signal-processing stages. One stage changes the gain on the detector's input; another increases uncertainty about the stimulus, allowing more noise into the detector; and other stages inject noise into the detector in a stimulus-dependent or -independent manner. By quantifying intrasaccadic suppression of flashed horizontal gratings at varying external noise levels, we obtained threshold-versus-noise (TVN) data, allowing us to fit the PTM. We tested if any of the PTM parameters changed significantly between the fixation and saccade models and could therefore account for intrasaccadic suppression. We found that the dominant contribution to intrasaccadic suppression was a reduction in the gain of the visual detector. We discuss how our study differs from previous ones that have pointed to uncertainty as an underlying cause of intrasaccadic suppression and how the equivalent noise approach provides a framework for comparing the disparate neural correlates of saccadic suppression.

An equivalent noise investigation of saccadic suppression.

Visual stimuli presented just before or during an eye movement are more difficult to detect than those same visual stimuli presented during fixation. This laboratory phenomenon--behavioral saccadic suppression--is thought to underlie the everyday experience of not perceiving the motion created by our own eye movements-saccadic omission. At the neural level, many cortical and subcortical areas respond differently to perisaccadic visual stimuli than to stimuli presented during fixation. Those neural response changes, however, are complex and the link to the behavioral phenomena of reduced detectability remains tentative. We used a well established model of human visual detection performance to provide a quantitative description of behavioral saccadic suppression and thereby allow a more focused search for its neural mechanisms. We used an equivalent noise method to distinguish between three mechanisms that could underlie saccadic suppression. The first hypothesized mechanism reduces the gain of the visual system, the second increases internal noise levels in a stimulus-dependent manner, and the third increases stimulus uncertainty. All three mechanisms predict that perisaccadic stimuli should be more difficult to detect, but each mechanism predicts a unique pattern of detectability as a function of the amount of external noise. Our experimental finding was that saccades increased detection thresholds at low external noise, but had little influence on thresholds at high levels of external noise. A formal analysis of these data in the equivalent noise analysis framework showed that the most parsimonious mechanism underlying saccadic suppression is a stimulus-independent reduction in response gain.

Receptive field positions in area MT during slow eye movements.

Perceptual stability requires the integration of information across eye movements. We first tested the hypothesis that motion signals are integrated by neurons whose receptive fields (RFs) do not move with the eye but stay fixed in the world. Specifically, we measured the RF properties of neurons in the middle temporal area (MT) of macaques (Macaca mulatta) during the slow phase of optokinetic nystagmus. Using a novel method to estimate RF locations for both spikes and local field potentials, we found that the location on the retina that changed spike rates or local field potentials did not change with eye position; RFs moved with the eye. Second, we tested the hypothesis that neurons link information across eye positions by remapping the retinal location of their RFs to future locations. To test this, we compared RF locations during leftward and rightward slow phases of optokinetic nystagmus. We found no evidence for remapping during slow eye movements; the RF location was not affected by eye-movement direction. Together, our results show that RFs of MT neurons and the aggregate activity reflected in local field potentials are yoked to the eye during slow eye movements. This implies that individual MT neurons do not integrate sensory information from a single position in the world across eye movements. Future research will have to determine whether such integration, and the construction of perceptual stability, takes place in the form of a distributed population code in eye-centered visual cortex or is deferred to downstream areas.

Transcranial electrical stimulation over visual cortex evokes phosphenes with a retinal origin.

Transcranial electrical stimulation (tES) is a promising therapeutic tool for a range of neurological diseases. Understanding how the small currents used in tES spread across the scalp and penetrate the brain will be important for the rational design of tES therapies. Alternating currents applied transcranially above visual cortex induce the perception of flashes of light (phosphenes). This makes the visual system a useful model to study tES. One hypothesis is that tES generates phosphenes by direct stimulation of the cortex underneath the transcranial electrode. Here, we provide evidence for the alternative hypothesis that phosphenes are generated in the retina by current spread from the occipital electrode. Building on the existing literature, we first confirm that phosphenes are induced at lower currents when electrodes are placed farther away from visual cortex and closer to the eye. Second, we explain the temporal frequency tuning of phosphenes based on the well-known response properties of primate retinal ganglion cells. Third, we show that there is no difference in the time it takes to evoke phosphenes in the retina or by stimulation above visual cortex. Together, these findings suggest that phosphenes induced by tES over visual cortex originate in the retina. From this, we infer that tES currents spread well beyond the area of stimulation and are unlikely to lead to focal neural activation. Novel stimulation protocols that optimize current distributions are needed to overcome these limitations of tES.

Steady state visual evoked potentials in schizophrenia: A review.

Over the past decades, researchers have explored altered rhythmic responses to visual stimulation in people with schizophrenia using steady state visual evoked potentials (SSVEPs). Here we systematically review studies performed between 1954 and 2021, as identified on PubMed. We included studies if they included people with schizophrenia, a control group, reported SSVEPs as their primary outcome, and used quantitative analyses in the frequency domain. We excluded studies that used SSVEPs to primarily quantify cognitive processes (e.g., attention). Fifteen studies met these criteria. These studies reported decreased SSVEPs across a range of frequencies and electrode locations in people living with schizophrenia compared to controls; none reported increases. Null results, however, were common. Given the typically modest number of subjects in these studies, this is consistent with a moderate effect size. It is notable that most studies targeted frequencies that fall within the alpha and beta band, and investigations of frequencies in the gamma band have been rare. We group test frequencies in frequency bands and summarize the results in topographic plots. From the wide range of approaches in these studies, we distill suggested experimental designs and analysis choices for future experiments. This will increase the value of SSVEP studies, improve our understanding of the mechanisms that result in altered rhythmic responses to visual stimulation in schizophrenia, and potentially further the development of diagnostic tools.

A checklist for assessing the methodological quality of concurrent tES-fMRI studies (ContES checklist): a consensus study and statement.

Low-intensity transcranial electrical stimulation (tES), including alternating or direct current stimulation, applies weak electrical stimulation to modulate the activity of brain circuits. Integration of tES with concurrent functional MRI (fMRI) allows for the mapping of neural activity during neuromodulation, supporting causal studies of both brain function and tES effects. Methodological aspects of tES-fMRI studies underpin the results, and reporting them in appropriate detail is required for reproducibility and interpretability. Despite the growing number of published reports, there are no consensus-based checklists for disclosing methodological details of concurrent tES-fMRI studies. The objective of this work was to develop a consensus-based checklist of reporting standards for concurrent tES-fMRI studies to support methodological rigor, transparency and reproducibility (ContES checklist). A two-phase Delphi consensus process was conducted by a steering committee (SC) of 13 members and 49 expert panelists through the International Network of the tES-fMRI Consortium. The process began with a circulation of a preliminary checklist of essential items and additional recommendations, developed by the SC on the basis of a systematic review of 57 concurrent tES-fMRI studies. Contributors were then invited to suggest revisions or additions to the initial checklist. After the revision phase, contributors rated the importance of the 17 essential items and 42 additional recommendations in the final checklist. The state of methodological transparency within the 57 reviewed concurrent tES-fMRI studies was then assessed by using the checklist. Experts refined the checklist through the revision and rating phases, leading to a checklist with three categories of essential items and additional recommendations: (i) technological factors, (ii) safety and noise tests and (iii) methodological factors. The level of reporting of checklist items varied among the 57 concurrent tES-fMRI papers, ranging from 24% to 76%. On average, 53% of checklist items were reported in a given article. In conclusion, use of the ContES checklist is expected to enhance the methodological reporting quality of future concurrent tES-fMRI studies and increase methodological transparency and reproducibility.

Summation of visual motion across eye movements reflects a nonspatial decision mechanism.

Human vision remains perceptually stable even though retinal inputs change rapidly with each eye movement. Although the neural basis of visual stability remains unknown, a recent psychophysical study pointed to the existence of visual feature-representations anchored in environmental rather than retinal coordinates (e.g., "spatiotopic" receptive fields; Melcher and Morrone, 2003). In that study, sensitivity to a moving stimulus presented after a saccadic eye movement was enhanced when preceded by another moving stimulus at the same spatial location before the saccade. The finding is consistent with spatiotopic sensory integration, but it could also have arisen from a probabilistic improvement in performance due to the presence of more than one motion signal for the perceptual decision. Here we show that this statistical advantage accounts completely for summation effects in this task. We first demonstrate that measurements of summation are confounded by noise related to an observer's uncertainty about motion onset times. When this uncertainty is minimized, comparable summation is observed regardless of whether two motion signals occupy the same or different locations in space, and whether they contain the same or opposite directions of motion. These results are incompatible with the tuning properties of motion-sensitive sensory neurons and provide no evidence for a spatiotopic representation of visual motion. Instead, summation in this context reflects a decision mechanism that uses abstract representations of sensory events to optimize choice behavior.

Weighted integration of visual position information.

The ability to localize visual objects is a fundamental component of human behavior and requires the integration of position information from object components. The retinal eccentricity of a stimulus and the locus of spatial attention can affect object localization, but it is unclear whether these factors alter the global localization of the object, the localization of object components, or both. We used psychophysical methods in humans to quantify behavioral responses in a centroid estimation task. Subjects located the centroid of briefly presented random dot patterns (RDPs). A peripheral cue was used to bias attention toward one side of the display. We found that although subjects were able to localize centroid positions reliably, they typically had a bias toward the fovea and a shift toward the locus of attention. We compared quantitative models that explain these effects either as biased global localization of the RDPs or as anisotropic integration of weighted dot component positions. A model that allowed retinal eccentricity and spatial attention to alter the weights assigned to individual dot positions best explained subjects' performance. These results show that global position perception depends on both the retinal eccentricity of stimulus components and their positions relative to the current locus of attention.