Non-Human Primate Eye-Tracking Publications
All EyeLink eye tracker non-human primate research publications up until 2025 (with some early 2026s) are listed below by year. You can search the eye-tracking publications using keywords such as Temporal Cortex, Macaque, Antisaccade, etc. You can also search for individual author names. If we missed any EyeLink non-human primate articles, please email us!
2008 |
Hans P. Op De Beeck; Jennifer A. Deutsch; Wim Vanduffel; Nancy Kanwisher; James J. DiCarlo A stable topography of selectivity for unfamiliar shape classes in monkey inferior temporal cortex Journal Article In: Cerebral Cortex, vol. 18, no. 7, pp. 1676–1694, 2008. @article{OpDeBeeck2008,The inferior temporal (IT) cortex in monkeys plays a central role in visual object recognition and learning. Previous studies have observed patches in IT cortex with strong selectivity for highly familiar object classes (e.g., faces), but the principles behind this functional organization are largely unknown due to the many properties that distinguish different object classes. To unconfound shape from meaning and memory, we scanned monkeys with functional magnetic resonance imaging while they viewed classes of initially novel objects. Our data revealed a topography of selectivity for these novel object classes across IT cortex. We found that this selectivity topography was highly reproducible and remarkably stable across a 3-month interval during which monkeys were extensively trained to discriminate among exemplars within one of the object classes. Furthermore, this selectivity topography was largely unaffected by changes in behavioral task and object retinal position, both of which preserve shape. In contrast, it was strongly influenced by changes in object shape. The topography was partially related to, but not explained by, the previously described pattern of face selectivity. Together, these results suggest that IT cortex contains a large-scale map of shape that is largely independent of meaning, familiarity, and behavioral task. |
Xiaochuan Pan; Kosuke Sawa; Ichiro Tsuda; Minoru Tsukada; Masamichi Sakagami Reward prediction based on stimulus categorization in primate lateral prefrontal cortex Journal Article In: Nature Neuroscience, vol. 11, no. 6, pp. 703–712, 2008. @article{Pan2008,To adapt to changeable or unfamiliar environments, it is important that animals develop strategies for goal-directed behaviors that meet the new challenges. We used a sequential paired-association task with asymmetric reward schedule to investigate how prefrontal neurons integrate multiple already-acquired associations to predict reward. Two types of reward-related neurons were observed in the lateral prefrontal cortex: one type predicted reward independent of physical properties of visual stimuli and the other encoded the reward value specific to a category of stimuli defined by the task requirements. Neurons of the latter type were able to predict reward on the basis of stimuli that had not yet been associated with reward, provided that another stimulus from the same category was paired with reward. The results suggest that prefrontal neurons can represent reward information on the basis of category and propagate this information to category members that have not been linked directly with any experience of reward. |
P. D. Thiem; Jessica A. Hill; K. -M. Lee; Edward L. Keller Behavioral properties of saccades generated as a choice response Journal Article In: Experimental Brain Research, vol. 186, no. 3, pp. 355–364, 2008. @article{Thiem2008,The behavior characterizing choice response decision-making was studied in monkeys to provide background information for ongoing neurophysiological studies of the neural mechanisms underlying saccadic choice decisions. Animals were trained to associate a specific color from a set of colored visual stimuli with a specific spatial location. The visual stimuli (colored disks) appeared briefly at equal eccentricity from a central fixation position and then were masked by gray disks. The correct target association was subsequently cued by the appearance of a colored stimulus at the fixation point. The animal indicated its choice by saccading to the remembered location of the eccentric stimulus, which had matched the color of the cue. The number of alternative associations (NA) varied from 1 to 4 and remained fixed within a block of trials. After the training period, performance (percent correct responses) declined modestly as NA increased (on average 96, 93 or 84% correct for 1, 2 or 4 NA, respectively). Response latency increased logarithmically as a function of NA, thus obeying Hick's law. The spatial extent of the learned association between color and location was investigated by rotating the array of colored stimuli that had remained fixed during the learning phase to various different angles. Error rates in choice saccades increased gradually as a function of the amount of rotation. The learned association biased the direction of the saccadic response toward the quadrant associated with the cue, but saccade direction was always toward one of the actual visual stimuli. This suggests that the learned associations between stimuli and responses were not spatially exact, but instead the association between color and location was distributed with declining strength from the trained locations. These results demonstrate that the saccade system in monkeys also displays the characteristic dependence on NA in choice response latencies, while more basic features of the eye movements are invariant from those in other tasks. The findings also provide behavioral evidence that spatially distributed regions are established for the sensory-to-motor associations during training which are later utilized for choice decisions. |
2007 |
Xiaodong Chen; Feng Han; Mu-ming Poo; Yang Dan Excitatory and suppressive receptive field subunits in awake monkey primary visual cortex (V1) Journal Article In: Proceedings of the National Academy of Sciences, vol. 104, no. 48, pp. 19120–19125, 2007. @article{Chen2007a,An essential step in understanding visual processing is to characterize the neuronal receptive fields (RFs) at each stage of the visual pathway. However, RF characterization beyond simple cells in the primary visual cortex (V1) remains a major challenge. Recent application of spike-triggered covariance (STC) analysis has greatly facilitated characterization of complex cell RFs in anesthetized animals. Here we apply STC to RF characterization in awake monkey V1. We found up to nine subunits for each cell, including one or two dominant excitatory subunits as described by the standard model, along with additional excitatory and suppressive subunits with weaker contributions. Compared with the dominant subunits, the nondominant excitatory subunits prefer similar orientations and spatial frequencies but have larger spatial envelopes. They contribute to response invariance to small changes in stimulus orientation, position, and spatial frequency. In contrast, the suppressive subunits are tuned to orientations 45 degrees -90 degrees different from the excitatory subunits, which may underlie cross-orientation suppression. Together, the excitatory and suppressive subunits form a compact description of RFs in awake monkey V1, allowing prediction of the responses to arbitrary visual stimuli. |
Benjamin Y. Hayden; Michael L. Platt Temporal discounting predicts risk sensitivity in rhesus macaques Journal Article In: Current Biology, vol. 17, no. 1, pp. 49–53, 2007. @article{Hayden2007,Humans and animals tend both to avoid uncertainty and to prefer immediate over future rewards. The comorbidity of psychiatric disorders such as impulsivity, problem gambling, and addiction suggests that a common mechanism may underlie risk sensitivity and temporal discounting [1-6]. Nonetheless, the precise relationship between these two traits remains largely unknown [3, 5]. To examine whether risk sensitivity and temporal discounting reflect a common process, we recorded choices made by two rhesus macaques in a visual gambling task [7] while we varied the delay between trials. We found that preference for the risky option declined with increasing delay between sequential choices in the task, even when all other task parameters were held constant. These results were quantitatively predicted by a model that assumed that the subjective expected utility of the risky option is evaluated based on the expected time of the larger payoff [5, 6]. The importance of the larger payoff in this model suggests that the salience of larger payoffs played a critical role in determining the value of risky options. These data suggest that risk sensitivity may be a product of other cognitive processes, and specifically that myopia for the future and the salience of jackpots control the propensity to take a gamble. |
Ryan E. B. Mruczek; David L. Sheinberg Activity of inferior temporal cortical neurons predicts recognition choice behavior and recognition time during visual search Journal Article In: Journal of Neuroscience, vol. 27, no. 11, pp. 2825–2836, 2007. @article{Mruczek2007,Although the selectivity for complex stimuli exhibited by neurons in inferior temporal cortex is often taken as evidence of their role in visual perception, few studies have directly tested this hypothesis. Here, we sought to create a relatively natural task with few behavioral constraints to test whether activity in inferior temporal cortex neurons predicts whether or not a monkey will recognize and respond to a complex visual object. Monkeys were trained to freely view an array of images and report the presence of one of many possible target images previously associated with a hand response. On certain trials, the identity of the target was swapped during the monkeys' targeting saccade. Furthermore, the response association of the preswap target and the postswap target differed (e.g., right-to-left target swap). Neural activity in cells selective for the preswap target was significantly higher when the monkeys' response matched the hand association of the preswap target. Furthermore, the monkeys' response time was predicted by the magnitude of the presaccadic firing rate on nonswap trials. Our results provide additional support for the role of inferior temporal cortex in object recognition during natural behavior. |
Ryan E. B. Mruczek; David L. Sheinberg Context familiarity enhances target processing by inferior temporal cortex neurons Journal Article In: Journal of Neuroscience, vol. 27, no. 32, pp. 8533–8545, 2007. @article{Mruczek2007a,Experience-dependent changes in the response properties of ventral visual stream neurons are thought to underlie our ability to rapidly and efficiently recognize visual objects. How these neural changes are related to efficient visual processing during natural vision remains unclear. Here, we demonstrate a neurophysiological correlate of efficient visual search through highly familiar object arrays. Humans and monkeys are faster at locating the same target when it is surrounded by familiar compared with unfamiliar distractors. We show that this behavioral enhancement is driven by an increased sensitivity of target-selective neurons in inferior temporal cortex. This results from an increased "signal" for target representations and decreased "noise" from neighboring familiar distractors. These data highlight the dynamic properties of the inferior temporal cortex neurons and add to a growing body of evidence demonstrating how experience shapes neural processing in the ventral visual stream. |
Zhi-Ming Shen; Wei-Feng Xu; Chao-Yi Li Cue-invariant detection of centre-surround discontinuity by V1 neurons in awake macaque monkey Journal Article In: Journal of Physiology, vol. 583, no. 2, pp. 581–592, 2007. @article{shen2007,Visual perception of an object depends on the discontinuity between the object and its background, which can be defined by a variety of visual features, such as luminance, colour and motion. While human object perception is largely cue invariant, the extent to which neural mechanisms in the primary visual cortex contribute to cue-invariant perception has not been examined extensively. Here we report that many V1 neurons in the awake monkey are sensitive to the stimulus discontinuity between their classical receptive field (CRF) and non-classical receptive field (nCRF) regardless of the visual feature that defines the discontinuity. The magnitude of this sensitivity is strongly dependent on the strength of nCRF suppression of the cell. These properties of V1 neurons may contribute significantly to cue-invariant object perception. |
Neil W. D. Thomas; Martin Pare Temporal processing of saccade targets in parietal cortex area LIP during visual search Journal Article In: Journal of Neurophysiology, vol. 97, no. 1, pp. 942–947, 2007. @article{Thomas2007a,We studied whether the lateral intraparietal (LIP) area—a subdivision of parietal cortex anatomically interposed between visual cortical areas and saccade executive centers—contains neurons with activity patterns sufficient to contribute to the active process of selecting saccade targets in visual search. Visually responsive neurons were recorded while monkeys searched for a color-different target presented concurrently with seven distractors evenly distributed in a circular search array. We found that LIP neurons initially responded indiscriminately to the presentation of a visual stimulus in their response fields, regardless of its feature and identity. Their activation nevertheless evolved to signal the search target before saccade initiation: an ideal observer could reliably discriminate the target from the individual activation of 60% of neurons, on average, 138 ms after stimulus presentation and 26 ms before saccade initiation. Importantly, the timing of LIP neuronal discrimination varied proportionally with reaction times. These findings suggest that LIP activity reflects the selection of both the search target and the targeting saccade during active visual search. |
2006 |
S. V. David; Benjamin Y. Hayden; Jack L. Gallant Spectral receptive field properties explain shape selectivity in area V4 Journal Article In: Journal of Neurophysiology, vol. 96, no. 6, pp. 3492–3505, 2006. @article{David2006,Neurons in cortical area V4 respond selectively to complex visual patterns such as curved contours and non-Cartesian gratings. Most previous experiments in V4 have measured responses to small, idiosyncratic stimulus sets and no single functional model yet accounts for all of the disparate results. We propose that one model, the spectral receptive field (SRF), can explain many observations of selectivity in V4. The SRF describes tuning in terms of the orientation and spatial frequency spectrum and can, in principle, predict the response to any visual stimulus. We estimated SRFs for neurons in V4 of awake primates by linearized reverse correlation of responses to a large set of natural images. We find that V4 neurons have large orientation and spatial frequency bandwidth and often bimodal orientation tuning. For comparison, we estimated SRFs for neurons in primary visual cortex (V1). Consistent with previous observations, we find that V1 neurons have narrower bandwidth than that of V4. To determine whether estimated SRFs can account for previous observations of selectivity, we used them to predict responses to Cartesian gratings, non-Cartesian gratings, natural images, and curved contours. Based on these predictions, we find that the majority of neurons in V1 are selective for Cartesian gratings, whereas the majority of V4 neurons are selective for non-Cartesian gratings or natural images. The SRF describes visual tuning properties with a second-order nonlinear model. These results support the hypothesis that a second-order model is sufficient to describe the general mechanisms mediating shape selectivity in area V4. |
Sarah J. Fairhall; Carol A. Dickson; Leah Scott; Peter C. Pearce A non-invasive method for studying an index of pupil diameter and visual performance in the rhesus monkey Journal Article In: Journal of Medical Primatology, vol. 35, no. 2, pp. 67–77, 2006. @article{Fairhall2006,BACKGROUND: A non-invasive model has been developed to estimate gaze direction and relative pupil diameter, in minimally restrained rhesus monkeys, to investigate the effects of low doses of ocularly administered cholinergic compounds on visual performance. METHODS: Animals were trained to co-operate with a novel device, which enabled eye movements to be recorded using modified human eye-tracking equipment, and to perform a task which determined visual threshold contrast. Responses were made by gaze transfer under twilight conditions. 4% w/v pilocarpine nitrate was studied to demonstrate the suitability of the model. RESULTS: Pilocarpine induced marked miosis for >3 h which was accompanied by a decrement in task performance. CONCLUSIONS: The method obviates the need for invasive surgery and, as the position of point of gaze can be approximately defined, the approach may have utility in other areas of research involving non-human primates. |
Kelly Shen; Martin Paré Guidance of eye movements during visual conjunction search: Local and global contextual effects on target discriminability Journal Article In: Journal of Neurophysiology, vol. 95, no. 5, pp. 2845–2855, 2006. @article{shen2006guidance,The composition of a visual scene influences the ability of humans to select specific details within that scene for discrimination or foveation with saccadic eye movements. With the goal of establishing an animal model to investigate the neural mechanisms underlying the deployment of visual attention and the guidance of saccades during visual search, we studied the visual behavior of three monkeys while they performed a conjunction (color + form) search task similar to those used in human studies. We found that search performance declined when distractors adjacent to the target shared its color, thereby revealing that color was more discriminable than form in these displays and suggesting that monkeys perceptually grouped stimuli by proximity and similarity. Search performance also varied with the overall composition of the display. Most importantly, saccades were biased toward distractors sharing the target color when there were few of them within the display and away from those distractors when they were numerous. Last, the monkeys initiated saccades with a fixed latency, suggesting that their responses to the display were automatic and that search strategies did not involve attentional resources beyond those recruited for regulating saccades. We conclude that monkeys adapt their visual strategies, largely via bottom-up processes, to both the local and the global context of the search. These findings suggest that the visual behavior of monkeys is guided by strategies similar to those observed in humans. |
2005 |
Benjamin Y. Hayden; Jack L. Gallant Time course of attention reveals different mechanisms for spatial and feature-based attention in area V4 Journal Article In: Neuron, vol. 47, no. 5, pp. 637–643, 2005. @article{Hayden2005,Attention can facilitate visual processing, emphasizing specific locations and highlighting stimuli containing specific features. To dissociate the mechanisms of spatial and feature-based attention, we compared the time course of visually evoked responses under different attention conditions. We recorded from single neurons in area V4 during a delayed match-to-sample task that controlled both spatial and feature-based attention. Neuronal responses increased when spatial attention was directed toward the receptive field and were modulated by the identity of the target of feature-based attention. Modulation by spatial attention was weaker during the early portion of the visual response and stronger during the later portion of the response. In contrast, modulation by feature-based attention was relatively constant throughout the response. It appears that stimulus onset transients disrupt spatial attention, but not feature attention. We conclude that spatial attention reflects a combination of stimulus-driven and goal-driven processes, while feature-based attention is purely goal driven. |
D. Zoccolan Multiple object response normalization in monkey inferotemporal cortex Journal Article In: Journal of Neuroscience, vol. 25, no. 36, pp. 8150–8164, 2005. @article{Zoccolan2005,The highest stages of the visual ventral pathway are commonly assumed to provide robust representation of object identity by disregard- ingconfoundingfactorssuchas object position, size, illumination,andthe presence of other objects (clutter).However,whereasneuronal responses in monkey inferotemporal cortex (IT) can show robust tolerance to position and size changes, previous work shows that responses to preferred objects are usually reduced by the presence of nonpreferred objects. More broadly, we do not yet understand multiple object representation in IT. In this study, we systematically examined IT responses to pairs and triplets of objects in three passively viewing monkeys across a broad range of object effectiveness.Wefound that, at least under these limited clutter conditions, a large fraction of the response of each IT neuron to multiple objects is reliably predicted as the average of its responses to the constituent objects in isolation. That is, multiple object responses depend primarilyonthe relative effectiveness of the constituent objects, regardless of object identity. This average effect becomes virtually perfect when populations of IT neurons are pooled. Furthermore, the average effect cannot simply be explained by attentional shifts but behaves as a primarily feedforward response property. Together, our obser- vations are most consistent with mechanistic models in which IT neuronal outputs are normalized by summed synaptic drive into IT or spiking activity within IT and suggest that normalization mechanisms previously revealed at earlier visual areas are operating through- out the ventral visual stream. |
2003 |
James A. Mazer; Jack L. Gallant Goal-related activity in V4 during free viewing visual search: Evidence for a ventral stream visual salience map Journal Article In: Neuron, vol. 40, no. 6, pp. 1241–1250, 2003. @article{Mazer2003,Natural exploration of complex visual scenes depends on saccadic eye movements toward important locations. Saccade targeting is thought to be mediated by a retinotopic map that represents the locations of salient features. In this report, we demonstrate that extrastriate ventral area V4 contains a retinotopic salience map that guides exploratory eye movements during a naturalistic free viewing visual search task. In more than half of recorded cells, visually driven activity is enhanced prior to saccades that move the fovea toward the location previously occupied by a neuron's spatial receptive field. This correlation suggests that bottom-up processing in V4 influences the oculomotor planning process. Half of the neurons also exhibit top-down modulation of visual responses that depends on search target identity but not visual stimulation. Convergence of bottom-up and top-down processing streams in area V4 results in an adaptive, dynamic map of salience that guides oculomotor planning during natural vision. |
2002 |
David A. Leopold; Holger K. Plettenberg; Nikos K. Logothetis Visual processing in the ketamine-anesthetized monkey: Optokinetic and blood oxygenation level-dependent responses Journal Article In: Experimental Brain Research, vol. 143, no. 3, pp. 359–372, 2002. @article{Leopold2002,We used optokinetic responses and functional magnetic resonance imaging (fMRI) to examine visual processing in monkeys whose conscious state was modulated by low doses (1-2 mg/kg) of the dissociative anesthetic ketamine. We found that, despite the animal's dissociated state and despite specific influences of ketamine on the oculomotor system, optokinetic nystagmus (OKN) could be reliably elicited with large, moving visual patterns. Responses were horizontally bidirectional for monocular stimulation, indicating that ketamine did not eliminate cortical processing of the motion stimulus. Also, results from fMRI directly demonstrated that the cortical blood oxygenation level-dependent (BOLD) response to visual patterns was preserved at the same ketamine doses used to elicit OKN. Finally, in the ketamine-anesthetized state, perceptually bistable motion stimuli produced patterns of spontaneously alternating OKN that normally would be tightly coupled to perceptual changes. These results, taken together, demonstrate that after ketamine administration cortical circuits continue to processes visual patterns in a dose-dependent manner despite the animal's behavioral dissociation. While perceptual experience is difficult to evaluate under these conditions, oculomotor patterns revealed that the brain not only registers but also acts upon its sensory input, employing it to drive a sensorimotor loop and even responding to a sensory conflict by engaging in spontaneous perception-related state changes. The ketamine-anesthetized monkey preparation thereby offers a safe and viable paradigm for the behavioral and electrophysiological investigation of issues related to conscious perception and anesthesia, as well as neural mechanisms of basic sensory processing. |