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1.  Optimal Control of Saccades by Spatial-Temporal Activity Patterns in the Monkey Superior Colliculus 
PLoS Computational Biology  2012;8(5):e1002508.
A major challenge in computational neurobiology is to understand how populations of noisy, broadly-tuned neurons produce accurate goal-directed actions such as saccades. Saccades are high-velocity eye movements that have stereotyped, nonlinear kinematics; their duration increases with amplitude, while peak eye-velocity saturates for large saccades. Recent theories suggest that these characteristics reflect a deliberate strategy that optimizes a speed-accuracy tradeoff in the presence of signal-dependent noise in the neural control signals. Here we argue that the midbrain superior colliculus (SC), a key sensorimotor interface that contains a topographically-organized map of saccade vectors, is in an ideal position to implement such an optimization principle. Most models attribute the nonlinear saccade kinematics to saturation in the brainstem pulse generator downstream from the SC. However, there is little data to support this assumption. We now present new neurophysiological evidence for an alternative scheme, which proposes that these properties reside in the spatial-temporal dynamics of SC activity. As predicted by this scheme, we found a remarkably systematic organization in the burst properties of saccade-related neurons along the rostral-to-caudal (i.e., amplitude-coding) dimension of the SC motor map: peak firing-rates systematically decrease for cells encoding larger saccades, while burst durations and skewness increase, suggesting that this spatial gradient underlies the increase in duration and skewness of the eye velocity profiles with amplitude. We also show that all neurons in the recruited population synchronize their burst profiles, indicating that the burst-timing of each cell is determined by the planned saccade vector in which it participates, rather than by its anatomical location. Together with the observation that saccade-related SC cells indeed show signal-dependent noise, this precisely tuned organization of SC burst activity strongly supports the notion of an optimal motor-control principle embedded in the SC motor map as it fully accounts for the straight trajectories and kinematic nonlinearity of saccades.
Author Summary
As the fovea is the only spot on the retina with high spatial resolution, primates need to move their eyes to peripheral targets for detailed inspection. Saccades are the fastest movements of the body, and theoretical studies suggest that their trajectories are optimized to bring the fovea as fast and accurately as possible on target. Speed-accuracy optimization principles explain the stereotyped nonlinear ‘main-sequence’ relationship between saccade amplitude, duration, and peak velocity. Earlier models attributed these kinematic properties to nonlinear neural circuitry in the brainstem but this creates problems for oblique saccades. Here, we demonstrate how the brainstem can be linear, and how instead the midbrain superior colliculus (SC) could optimize saccadic speed-accuracy tradeoff. Each saccade involves the recruitment of a large population of SC neurons. We show that peak firing-rate and burst shape of the recruited cells systematically vary with their location in the SC, and that burst shapes nicely match the eye-velocity profiles. This organization of burst properties fully explains the main-sequence. Moreover, all cells synchronize their bursts, thus maximizing the total instantaneous input to the brainstem, and ensuring that oblique saccades have straight trajectories. We thus discovered a sophisticated neural mechanism underlying optimal motor control in the brain.
PMCID: PMC3355059  PMID: 22615548
2.  Saccadic Adaptation to Moving Targets 
PLoS ONE  2012;7(6):e39708.
Saccades are so called ballistic movements which are executed without online visual feedback. After each saccade the saccadic motor plan is modified in response to post-saccadic feedback with the mechanism of saccadic adaptation. The post-saccadic feedback is provided by the retinal position of the target after the saccade. If the target moves after the saccade, gaze may follow the moving target. In that case, the eyes are controlled by the pursuit system, a system that controls smooth eye movements. Although these two systems have in the past been considered as mostly independent, recent lines of research point towards many interactions between them. We were interested in the question if saccade amplitude adaptation is induced when the target moves smoothly after the saccade. Prior studies of saccadic adaptation have considered intra-saccadic target steps as learning signals. In the present study, the intra-saccadic target step of the McLaughlin paradigm of saccadic adaptation was replaced by target movement, and a post-saccadic pursuit of the target. We found that saccadic adaptation occurred in this situation, a further indication of an interaction of the saccadic system and the pursuit system with the aim of optimized eye movements.
PMCID: PMC3387193  PMID: 22768109
3.  Infant eye and head movements toward the side opposite the cue in the anti-saccade paradigm 
The anti-saccade task, when people must respond in the direction opposite to a visual stimulus, has been used as a marker of operation of the frontal cortical oculomotor area. However, early development of oculomotor control has been little studied with the infant anti-saccade paradigm, and a few studies did not recognize anti-saccades in infants in light of the results of adult anti-saccade. Since the characteristics of infant eye movements are little known, applying the criteria used in adult study is by no means the best way to study infant anti-saccade. As it is indicated that coordinated eye and head movements often enable infants to control the direction of their gaze, head movements should be examined as an infant orienting response. The aim of this study was to address how infants used eye and head movements during the anti-saccade paradigm. To distinguish infants' responses, we also investigated eye and head movements during a task for an inhibition of return. Inhibition of return, in which delayed responses occur in the direction to which attention had previously been oriented, has been thought to mark activity of the superior colliculus. Since the superior colliculus is thought to develop much earlier in life than the frontal lobes, we thought it useful to compare these task performances during infancy.
Infants were divided into three groups according to age. Anti-saccade and inhibition-of-return tasks were given. Their eye and head movements during tasks were independently recorded by the corneal reflection method in the head-free condition.
Younger infants tended to initiate eye movement less than older ones in both tasks. In the anti-saccade task, responses opposite to the cue tended to show longer latency than responses to the cue. Infants made faster responses toward the side opposite the cue when it was to the right than when it was left of fixation. Regarding the comparison of responses toward the side opposite the cue between two tasks, the leftward eye movement was faster than the leftward head movements in the inhibition-of-return task, while no difference of latency was observed between eye and head movements in the anti-saccade task. A qualitative analysis of the trajectory of these responses revealed that head movement trajectories were steeper in the anti-saccade than in the inhibition-of-return task.
Younger infants move head and eyes together, with head movements frequently starting first. On the other hand, both the leftward latency difference between eye and head and gentle trajectories of head in inhibition of return indicate that eye movements are more predominant over head movements in the inhibition-of-return task than in the anti-saccade task. This would suggest an earlier developing inhibition-of-return mechanism.
PMCID: PMC1783854  PMID: 17229319
4.  Neurobehavioral Abnormalities in First-Degree Relatives of Individuals With Autism 
Archives of general psychiatry  2010;67(8):830-840.
Studying sensorimotor and neurocognitive impairments in unaffected family members of individuals with autism may help identify familial pathophysiological mechanisms associated with the disorder.
To determine whether atypical sensorimotor or neurocognitive characteristics associated with autism are present in first-degree relatives of individuals with autism.
Case-control comparison of neurobehavioral functions.
University medical center.
Fifty-seven first-degree relatives of individuals with autism and 40 age-, sex-, and IQ-matched healthy control participants (aged 8–54 years).
Main Outcome Measures
Oculomotor tests of sensorimotor responses (saccades and smooth pursuit); procedural learning and response inhibition; neuropsychological tests of motor, memory, and executive functions; and psychological measures of social behavior, communication skills, and obsessive-compulsive behaviors.
On eye movement testing, family members demonstrated saccadic hypometria, reduced steady-state pursuit gain, and a higher rate of voluntary response inhibition errors relative to controls. They also showed lateralized deficits in procedural learning and open-loop pursuit gain (initial 100 milliseconds of pursuit) and increased variability in the accuracy of large-amplitude saccades that were confined to rightward movements. In neuropsychological studies, only executive functions were impaired relative to those of controls. Family members reported more communication abnormalities and obsessive-compulsive behaviors than controls. Deficits across oculomotor, neuropsychological, and psychological domains were relatively independent from one another.
Family members of individuals with autism demonstrate oculomotor abnormalities implicating pontocerebellar and frontostriatal circuits and left-lateralized alterations of frontotemporal circuitry and striatum. The left-lateralized alterations have not been identified in other neuropsychiatric disorders and are of interest given atypical brain lateralization and language development associated with the disorder. Similar oculomotor deficits have been reported in individuals with autism, suggesting that they may be familial and useful for studies of neurophysiological and genetic mechanisms in autism.
PMCID: PMC3145411  PMID: 20679591
5.  TMS perturbs saccade trajectories and unmasks an internal feedback controller for saccades 
When we applied a single pulse of transcranial magnetic stimulation (TMS) to any part of the human head during a saccadic eye movement, the ongoing eye velocity was reduced starting as early as 45ms after the TMS, and lasted around 32ms. The perturbation to the saccade trajectory was not due to a mechanical effect of the lid on the eye (e.g., from blinks). When the saccade involved coordinated movements of both the eyes and the lids, e.g., in vertical saccades, TMS produced a synchronized inhibition of the motor commands to both eye and lid muscles. The TMS induced perturbation of the eye trajectory did not show habituation with repetition, and was present in both pro- and anti-saccades. Despite the perturbation, the eye trajectory was corrected within the same saccade with compensatory motor commands that guided the eyes to the target. This within-saccade correction did not rely on visual input, suggesting that the brain monitored the oculomotor commands as the saccade unfolded, maintained a real time estimate of the position of the eyes, and corrected for the perturbation. TMS disrupted saccades regardless of the location of the coil on the head, suggesting that the coil discharge engages a non-habituating startle-like reflex system. This system affects ongoing motor commands upstream of the oculomotor neurons, possibly at the level of the superior colliculus or omnipause neurons. Therefore, a TMS pulse centrally perturbs saccadic motor commands, which are monitored possibly via efference copy, and are corrected via internal feedback.
PMCID: PMC3167087  PMID: 21832184
saccade accuracy; pause; TMS; startle; omnipause neuron; forward model
6.  Visual working memory modulates low-level saccade target selection: Evidence from rapidly generated saccades in the global effect paradigm 
Journal of Vision  2013;13(13):4.
In three experiments, we examined the influence of visual working memory (VWM) on the metrics of saccade landing position in a global effect paradigm. Participants executed a saccade to the more eccentric object in an object pair appearing on the horizontal midline, to the left or right of central fixation. While completing the saccade task, participants maintained a color in VWM for an unrelated memory task. Either the color of the saccade target matched the memory color (target match), the color of the distractor matched the memory color (distractor match), or the colors of neither object matched the memory color (no match). In the no-match condition, saccades tended to land at the midpoint between the two objects: the global, or averaging, effect. However, when one of the two objects matched VWM, the distribution of landing position shifted toward the matching object, both for target match and for distractor match. VWM modulation of landing position was observed even for the fastest quartile of saccades, with a mean latency as low as 112 ms. Effects of VWM on such rapidly generated saccades, with latencies in the express-saccade range, indicate that VWM interacts with the initial sweep of visual sensory processing, modulating perceptual input to oculomotor systems and thereby biasing oculomotor selection. As a result, differences in memory match produce effects on landing position similar to the effects generated by differences in physical salience.
PMCID: PMC3817959  PMID: 24190909
saccadic eye movements; visual working memory; visual short-term memory
7.  Brain Processing of Visual Information during Fast Eye Movements Maintains Motor Performance 
PLoS ONE  2013;8(1):e54641.
Movement accuracy depends crucially on the ability to detect errors while actions are being performed. When inaccuracies occur repeatedly, both an immediate motor correction and a progressive adaptation of the motor command can unfold. Of all the movements in the motor repertoire of humans, saccadic eye movements are the fastest. Due to the high speed of saccades, and to the impairment of visual perception during saccades, a phenomenon called “saccadic suppression”, it is widely believed that the adaptive mechanisms maintaining saccadic performance depend critically on visual error signals acquired after saccade completion. Here, we demonstrate that, contrary to this widespread view, saccadic adaptation can be based entirely on visual information presented during saccades. Our results show that visual error signals introduced during saccade execution–by shifting a visual target at saccade onset and blanking it at saccade offset–induce the same level of adaptation as error signals, presented for the same duration, but after saccade completion. In addition, they reveal that this processing of intra-saccadic visual information for adaptation depends critically on visual information presented during the deceleration phase, but not the acceleration phase, of the saccade. These findings demonstrate that the human central nervous system can use short intra-saccadic glimpses of visual information for motor adaptation, and they call for a reappraisal of current models of saccadic adaptation.
PMCID: PMC3558515  PMID: 23382932
8.  fMRI Evidence for Multisensory Recruitment Associated With Rapid Eye Movements During Sleep 
Human brain mapping  2009;30(5):1705-1722.
We studied the neural correlates of rapid eye movement during sleep (REM) by timing REMs from video recording and using rapid event-related functional MRI. Consistent with the hypothesis that REMs share the brain systems and mechanisms with waking eye movements and are visually-targeted saccades, we found REM-locked activation in the primary visual cortex, thalamic reticular nucleus (TRN), ‘visual claustrum’, retrosplenial cortex (RSC, only on the right hemisphere), fusiform gyrus, anterior cingulate cortex, and the oculomotor circuit that controls awake saccadic eye movements (and subserves awake visuospatial attention). Unexpectedly, robust activation also occurred in non-visual sensory cortices, motor cortex, language areas, and the ascending reticular activating system, including basal forebrain, the major source of cholinergic input to the entire cortex. REM-associated activation of these areas, especially non-visual primary sensory cortices, TRN and claustrum, parallels findings from waking studies on the interactions between multiple sensory data, and their ‘binding’ into a unified percept, suggesting that these mechanisms are also shared in waking and dreaming and that the sharing goes beyond the expected visual scanning mechanisms. Surprisingly, REMs were associated with a decrease in signal in specific periventricular subregions, matching the distribution of the serotonergic supraependymal plexus. REMs might serve as a useful task-free probe into major brain systems for functional brain imaging.
PMCID: PMC2753360  PMID: 18972392
fMRI; REM; thalamic reticular nucleus; multisensory; oculomotor circuit; visual scanning
9.  Evidence for Optimal Integration of Visual Feature Representations across Saccades 
The Journal of Neuroscience  2015;35(28):10146-10153.
We explore the visual world through saccadic eye movements, but saccades also present a challenge to visual processing by shifting externally stable objects from one retinal location to another. The brain could solve this problem in two ways: by overwriting preceding input and starting afresh with each fixation or by maintaining a representation of presaccadic visual features in working memory and updating it with new information from the remapped location. Crucially, when multiple objects are present in a scene the planning of eye movements profoundly affects the precision of their working memory representations, transferring limited memory resources from fixation toward the saccade target. Here we show that when humans make saccades, it results in an update of not just the precision of representations but also their contents. When multiple item colors are shifted imperceptibly during a saccade the perceived colors are found to fall between presaccadic and postsaccadic values, with the weight given to each input varying continuously with item location, and fixed relative to saccade parameters. Increasing sensory uncertainty, by adding color noise, biases updating toward the more reliable input, which is consistent with an optimal integration of presaccadic working memory with a postsaccadic updating signal. We recover this update signal and show it to be tightly focused on the vicinity of the saccade target. These results reveal how the nervous system accumulates detailed visual information from multiple views of the same object or scene.
SIGNIFICANCE STATEMENT This study examines the consequences of saccadic eye movements for the internal representation of visual objects. A saccade shifts the image of a stable visual object from one part of the retina to another. We show that visual representations are built up over these different views of the same object, by combining information obtained before and after each saccade. The weights given to presaccadic and postsaccadic information are determined by the relative reliability of each input. This provides evidence that the visual system combines inputs over time in a statistically optimal way.
PMCID: PMC4502255  PMID: 26180191
optimal integration; saccades; visual updating; working memory
10.  Adaptive control of saccades via internal feedback 
Ballistic movements like saccades require the brain to generate motor commands without the benefit of sensory feedback. Despite this, saccades are remarkably accurate. Theory suggests that this accuracy arises because the brain relies on an internal forward model that monitors the motor commands, predicts their sensory consequences, and corrects eye trajectory midflight. If control of saccades relies on a forward model, then the forward model should adapt whenever its predictions fail to match sensory feedback at the end of the movement. Using optimal feedback control theory, we predicted how this adaptation should alter saccade trajectories. We trained subjects on a paradigm where the horizontal target jumped vertically during the saccade. With training, the final position of the saccade moved toward the second target. However, saccades became increasingly curved, i.e., suboptimal, as oculomotor commands were corrected online to steer the eye toward the second target. The adaptive response had two components: 1) the motor commands that initiated the saccades changed slowly, aiming the saccade closer to the jumped target. The adaptation of these earliest motor commands displayed little forgetting during the rest periods. 2) Late in saccade trajectory, another adaptive response steered it still closer to the jumped target, producing curvature. Adaptation of these late motor commands showed near complete forgetting during the rest periods. The two components adapted at different timescales, with the late-acting component displaying much faster rates. It appears that in controlling saccades, the brain relies on an internal feedback that has the characteristics of a fast adapting forward model.
PMCID: PMC2733833  PMID: 18337410
saccade adaptation; forward model; internal feedback; optimal control; curved saccades; fatigue
11.  Sensory Processing of Motor Inaccuracy Depends on Previously Performed Movement and on Subsequent Motor Corrections: A Study of the Saccadic System 
PLoS ONE  2011;6(2):e17329.
When goal-directed movements are inaccurate, two responses are generated by the brain: a fast motor correction toward the target and an adaptive motor recalibration developing progressively across subsequent trials. For the saccadic system, there is a clear dissociation between the fast motor correction (corrective saccade production) and the adaptive motor recalibration (primary saccade modification). Error signals used to trigger corrective saccades and to induce adaptation are based on post-saccadic visual feedback. The goal of this study was to determine if similar or different error signals are involved in saccadic adaptation and in corrective saccade generation. Saccadic accuracy was experimentally altered by systematically displacing the visual target during motor execution. Post-saccadic error signals were studied by manipulating visual information in two ways. First, the duration of the displaced target after primary saccade termination was set at 15, 50, 100 or 800 ms in different adaptation sessions. Second, in some sessions, the displaced target was followed by a visual mask that interfered with visual processing. Because they rely on different mechanisms, the adaptation of reactive saccades and the adaptation of voluntary saccades were both evaluated. We found that saccadic adaptation and corrective saccade production were both affected by the manipulations of post-saccadic visual information, but in different ways. This first finding suggests that different types of error signal processing are involved in the induction of these two motor corrections. Interestingly, voluntary saccades required a longer duration of post-saccadic target presentation to reach the same amount of adaptation as reactive saccades. Finally, the visual mask interfered with the production of corrective saccades only during the voluntary saccades adaptation task. These last observations suggest that post-saccadic perception depends on the previously performed action and that the differences between saccade categories of motor correction and adaptation occur at an early level of visual processing.
PMCID: PMC3044175  PMID: 21383849
12.  Individual Differences in Impulsivity Predict Anticipatory Eye Movements 
PLoS ONE  2011;6(10):e26699.
Impulsivity is the tendency to act without forethought. It is a personality trait commonly used in the diagnosis of many psychiatric diseases. In clinical practice, impulsivity is estimated using written questionnaires. However, answers to questions might be subject to personal biases and misinterpretations. In order to alleviate this problem, eye movements could be used to study differences in decision processes related to impulsivity. Therefore, we investigated correlations between impulsivity scores obtained with a questionnaire in healthy subjects and characteristics of their anticipatory eye movements in a simple smooth pursuit task. Healthy subjects were asked to answer the UPPS questionnaire (Urgency Premeditation Perseverance and Sensation seeking Impulsive Behavior scale), which distinguishes four independent dimensions of impulsivity: Urgency, lack of Premeditation, lack of Perseverance, and Sensation seeking. The same subjects took part in an oculomotor task that consisted of pursuing a target that moved in a predictable direction. This task reliably evoked anticipatory saccades and smooth eye movements. We found that eye movement characteristics such as latency and velocity were significantly correlated with UPPS scores. The specific correlations between distinct UPPS factors and oculomotor anticipation parameters support the validity of the UPPS construct and corroborate neurobiological explanations for impulsivity. We suggest that the oculomotor approach of impulsivity put forth in the present study could help bridge the gap between psychiatry and physiology.
PMCID: PMC3202566  PMID: 22046334
13.  Manifestation of ocular-muscle EMG contamination in human intracranial recordings 
NeuroImage  2010;54(1):213-233.
It is widely assumed that intracranial recordings from the brain are only minimally affected by contamination due to ocular-muscle electromyogram (oEMG). Here we show that this is not always the case. In intracranial recordings from five surgical epilepsy patients we observed that eye movements caused a transient biphasic potential at the onset of a saccade, resembling the saccadic spike potential commonly seen in scalp EEG, accompanied by an increase in broadband power between 20 and 200 Hz. Using concurrently recorded eye movements and high-density intracranial EEG (iEEG) we developed a detailed overview of the spatial distribution and temporal characteristics of the saccade-related oculomotor signal within recordings from ventral, medial and lateral temporal cortex. The occurrence of the saccadic spike was not explained solely by reference contact location, and was observed near the temporal pole for small (< 2 deg) amplitude saccades and over a broad area for larger saccades. We further examined the influence of saccade-related oEMG contamination on measurements of spectral power and interchannel coherence. Contamination manifested in both spectral power and coherence measurements, in particular, over the anterior half of the ventral and medial temporal lobe. Next, we compared methods for removing the contaminating signal and found that nearest-neighbor bipolar re-referencing and ICA filtering were effective for suppressing oEMG at locations far from the orbits, but tended to leave some residual contamination at the temporal pole. Finally, we show that genuine cortical broadband gamma responses observed in averaged data from ventral temporal cortex can bear a striking similarity in time course and band-width to oEMG contamination recorded at more anterior locations. We conclude that eye movement-related contamination should be ruled out when reporting high gamma responses in human intracranial recordings, especially those obtained near anterior and medial temporal lobe.
PMCID: PMC2975438  PMID: 20696256
intracranial EEG; ECoG; saccadic spike; EMG; gamma band; Eye Movement; Eye Muscle
14.  Temporal Production Signals in Parietal Cortex 
PLoS Biology  2012;10(10):e1001413.
Neuronal activity in the lateral intraparietal cortex of monkeys reflects the passage of time and is sufficiently precise to explain timed eye movements.
We often perform movements and actions on the basis of internal motivations and without any explicit instructions or cues. One common example of such behaviors is our ability to initiate movements solely on the basis of an internally generated sense of the passage of time. In order to isolate the neuronal signals responsible for such timed behaviors, we devised a task that requires nonhuman primates to move their eyes consistently at regular time intervals in the absence of any external stimulus events and without an immediate expectation of reward. Despite the lack of sensory information, we found that animals were remarkably precise and consistent in timed behaviors, with standard deviations on the order of 100 ms. To examine the potential neural basis of this precision, we recorded from single neurons in the lateral intraparietal area (LIP), which has been implicated in the planning and execution of eye movements. In contrast to previous studies that observed a build-up of activity associated with the passage of time, we found that LIP activity decreased at a constant rate between timed movements. Moreover, the magnitude of activity was predictive of the timing of the impending movement. Interestingly, this relationship depended on eye movement direction: activity was negatively correlated with timing when the upcoming saccade was toward the neuron's response field and positively correlated when the upcoming saccade was directed away from the response field. This suggests that LIP activity encodes timed movements in a push-pull manner by signaling for both saccade initiation towards one target and prolonged fixation for the other target. Thus timed movements in this task appear to reflect the competition between local populations of task relevant neurons rather than a global timing signal.
Author Summary
One advantage of studying sensory systems is that external stimuli can be readily controlled and quantified. Our internal perception of time, however, is not as easily approachable. To address this challenge, we developed a task that leverages our knowledge of the neuronal circuits underlying eye movements. Monkeys were trained to move their eyes consistently at regular time intervals without any external cueing or immediate expectation of reward. The animals were remarkably precise and consistent in their timing. Recordings from individual lateral intraparietal area (LIP) neurons, an area associated with eye movement planning, showed a linear decrease in activity between eye movements. This contrasts with previous studies that have reported increases in LIP activity immediately prior to eye movements and suggests that expectation of reward may have influenced such results. We also found that variations in this decreasing activity were predictive of the animals’ timing. Finally, we demonstrate through a simple model that LIP activity was sufficiently precise to completely explain the timing of the animals' actions. The model demonstrates that a precise internal sense of time can arise by comparing activity across different neuronal populations.
PMCID: PMC3484129  PMID: 23118614
15.  A step towards non-invasive characterization of the human frontal eye fields of individual subjects 
Nonlinear Biomedical Physics  2010;4(Suppl 1):S11.
Identifying eye movement related areas in the frontal lobe has a long history, with microstimulation in monkeys producing the most clear-cut results. For humans, however, there is still no consensus about the location and the extent of the frontal eye field (FEF). There is also no simple non-invasive method for unambiguously defining the FEF in individual subjects, a prerequisite for clinical applications. Here we explore the use of magnetoencephalography (MEG) for the non-invasive identification and characterization of FEF activity in an individual subject.
We mapped human brain activity before, during and after saccades by applying tomographic analysis to MEG data. Statistical parametric maps and circular statistics produced plausible FEF loci, but no unambiguous definition for individual subjects. Here we first computed the spectral decomposition and correlation with electrooculogram (EOG) of the tomographic brain activations. For each of these two measures statistical comparisons were made between different saccades.
In this paper, we first review the frontal cortex activations identified in earlier animal and human studies and place the putative human FEFs in a well-defined anatomical framework. This framework is then used as reference for describing the results of new Fourier analysis of the tomographic solutions comparing active saccade tasks and their controls. The most consistent change in the dorsal frontal cortex was at the putative left FEF, for both saccades to the left and right. The asymmetric result is consistent with the 1-way callosal traffic theory. We also showed that the new correlation analysis had its most consistent change in the contralateral putative FEF. This result was obtained for EOG latencies before saccade onset with delays of a few hundreds of milliseconds (FEF activity leading the EOG) and only for visual cues signaling the execution of a saccade in a previously defined saccade direction.
The FEF definition derived from microstimulation describes only one of the areas in the dorsal lateral frontal lobe that act together to plan, prepare and execute a saccade. The definition and characterization of these areas in an individual subject can be obtained from non-invasive MEG measurements.
PMCID: PMC2880797  PMID: 20522261
16.  Individual Differences in the Real-Time Comprehension of Children with ASD 
Lay Abstract
Spoken language processing is related to language and cognitive skills in typically developing children, but very little is known about how children with autism spectrum disorders (ASD) comprehend words in real time. Studying this area is important because it may help us understand why many children with autism have delayed language comprehension. Thirty-four children with ASD (3–6 years old) participated in this study. They took part in a language comprehension task that involved looking at pictures on a screen and listening to questions about familiar nouns (e.g., Where’s the shoe?). Children as a group understood the familiar words, but accuracy and processing speed varied considerably across children. The children who were more accurate were also faster to process the familiar words. Children’s language processing accuracy was related to processing speed and language comprehension on a standardized test; nonverbal cognition did not explain additional information after accounting for these factors. Additionally, lexical processing accuracy at age 5½ was related to children’s vocabulary comprehension three years earlier, at age 2½. Autism severity and years of maternal education were unrelated to language processing. Words typically acquired earlier in life were processed more quickly than words acquired later. These findings point to similarities in patterns of language development in typically developing children and children with ASD. Studying real-time comprehension in children with ASD may help us better understand mechanisms of language comprehension in this population. Future work may help explain why some children with ASD develop age-appropriate language skills, whereas others experience lasting deficits.
Scientific Abstract
Many children with autism spectrum disorders (ASD) demonstrate deficits in language comprehension, but little is known about how they process spoken language as it unfolds. Real-time lexical comprehension is associated with language and cognition in children without ASD, suggesting that this may also be the case for children with ASD. This study adopted an individual differences approach to characterizing real-time comprehension of familiar words in a group of 34 three- to six-year-olds with ASD. The looking-while-listening paradigm was employed; it measures online accuracy and latency through language-mediated eye movements and has limited task demands. On average, children demonstrated comprehension of the familiar words, but considerable variability emerged. Children with better accuracy were faster to process the familiar words. In combination, processing speed and comprehension on a standardized language assessment explained 63% of the variance in online accuracy. Online accuracy was not correlated with autism severity or maternal education, and nonverbal cognition did not explain unique variance. Notably, online accuracy at age 5½ was related to vocabulary comprehension three years earlier. The words typically learned earliest in life were processed most quickly. Consistent with a dimensional view of language abilities, these findings point to similarities in patterns of language acquisition in typically developing children and those with ASD. Overall, our results emphasize the value of examining individual differences in real-time language comprehension in this population. We propose that the looking-while-listening paradigm is a sensitive and valuable methodological tool that can be applied across many areas of autism research.
PMCID: PMC3808474  PMID: 23696214
autism; comprehension; language processing; receptive vocabulary; eye-gaze methodology; individual differences
17.  Second language proficiency modulates conflict-monitoring in an oculomotor Stroop task: evidence from Hindi-English bilinguals 
Many studies have confirmed the presence of a bilingual advantage which is manifested as enhanced cognitive and attention control. However, very few studies have investigated the role of second language proficiency on the modulation of conflict-monitoring in bilinguals. We investigated this by comparing high and low proficient Hindi-English bilinguals on a modified saccadic arrow Stroop task under different monitoring conditions, and tested the predictions of the bilingual executive control advantage proposal. The task of the participants was to make an eye movement toward the color patch in the same color as the central arrow, ignoring the patch to which the arrow was pointing. High-proficient bilinguals had overall faster saccade latency on all types of trials as compared to the low proficient bilinguals. The overall saccadic latency for high proficiency bilinguals was similarly affected by the different types of monitoring conditions, whereas conflict resolution advantage was found only for high monitoring demanding condition. The results support a conflict-monitoring account in a novel oculomotor task and also suggest that language proficiency could modulate executive control in bilinguals.
PMCID: PMC3679481  PMID: 23781210
bilingualism; conflict-monitoring; language proficiency; saccades; Stroop task
18.  Processing of Hand-Related Verbs Specifically Affects the Planning and Execution of Arm Reaching Movements 
PLoS ONE  2012;7(4):e35403.
Even though a growing body of research has shown that the processing of action language affects the planning and execution of motor acts, several aspects of this interaction are still hotly debated. The directionality (i.e. does understanding action-related language induce a facilitation or an interference with the corresponding action?), the time course, and the nature of the interaction (i.e. under what conditions does the phenomenon occur?) are largely unclear. To further explore this topic we exploited a go/no-go paradigm in which healthy participants were required to perform arm reaching movements toward a target when verbs expressing either hand or foot actions were shown, and to refrain from moving when abstract verbs were presented. We found that reaction times (RT) and percentages of errors increased when the verb involved the same effector used to give the response. This interference occurred very early, when the interval between verb presentation and the delivery of the go signal was 50 ms, and could be elicited until this delay was about 600 ms. In addition, RTs were faster when subjects used the right arm than when they used the left arm, suggesting that action–verb understanding is left-lateralized. Furthermore, when the color of the printed verb and not its meaning was the cue for movement execution the differences between RTs and error percentages between verb categories disappeared, unequivocally indicating that the phenomenon occurs only when the semantic content of a verb has to be retrieved. These results are compatible with the theory of embodied language, which hypothesizes that comprehending verbal descriptions of actions relies on an internal simulation of the sensory–motor experience of the action, and provide a new and detailed view of the interplay between action language and motor acts.
PMCID: PMC3335064  PMID: 22536380
19.  Performance Deficits in a Voluntary Saccade Task in Chinese “Express Saccade Makers” 
PLoS ONE  2012;7(10):e47688.
Differences in behaviour and cognition have been observed in different human populations. It has been reported that in various types of complex visual task, eye movement patterns differ systematically between Chinese and non-Chinese participants, an observation that has been related to differences in culture between groups. However, we confirm here that, in healthy, naïve adult Chinese participants, a far higher proportion (22%) than expected (1–5%) exhibit a pattern of reflexive eye movement behaviour (high numbers of low latency express saccades) in circumstances designed to inhibit such responses (prosaccade overlap tasks). These participants are defined as “express saccade makers” (ESMs). We then show using the antisaccade paradigm, which requires the inhibition of reflexive responses and the programming and execution of voluntary saccades, that the performance of ESMs is compromised; they have higher antisaccade directional error rates, and the latency distributions of their error saccades again exhibit a higher proportion of low latency express saccade errors consistent with a reduced ability to inhibit reflexive responses. These results are difficult to reconcile with a cultural explanation as they relate to important and specific performance differences within a particular population. They suggest a potential unexpected confound relevant to those studies of Chinese versus other groups which have investigated group differences using oculomotor measures, and explained them in terms of culture. The confirmation of higher numbers of ESMs among Chinese participants provides new opportunities for examining oculomotor control.
PMCID: PMC3472994  PMID: 23091639
20.  Language Structures Used by Kindergartners with Cochlear Implants: Relationship to Phonological Awareness, Lexical Knowledge and Hearing Loss 
Ear and hearing  2014;35(5):506-518.
Listeners use their knowledge of how language is structured to aid speech recognition in everyday communication. When it comes to children with congenital hearing loss severe enough to warrant cochlear implants (CIs), the question arises of whether these children can acquire the language knowledge needed to aid speech recognition, in spite of only having spectrally degraded signals available to them. That question was addressed in the current study. Specifically there were three goals: (1) to compare the language structures used by children with CIs to those of children with normal hearing (NH); (2) to assess the amount of variance in the language measures explained by phonological awareness and lexical knowledge; and (3) to assess the amount of variance in the language measures explained by factors related to the hearing loss itself and subsequent treatment.
Language samples were obtained and transcribed for 40 children who had just completed kindergarten: 19 with NH and 21 with CIs. Five measures were derived from Systematic Analysis of Language Transcripts (SALT): (1) mean length of utterance in morphemes, (2) number of conjunctions, excluding and, (3) number of personal pronouns, (4) number of bound morphemes, and (5) number of different words. Measures were also collected on phonological awareness and lexical knowledge. Statistics examined group differences, as well as the amount of variance in the language measures explained by phonological awareness, lexical knowledge, and factors related to hearing loss and its treatment for children with CIs.
Mean scores of children with CIs were roughly one standard deviation below those of children with NH on all language measures, including lexical knowledge, matching outcomes of other studies. Mean scores of children with CIs were closer to two standard deviations below those of children with NH on two out of three measures of phonological awareness (specifically those related to phonemic structure). Lexical knowledge explained significant amounts of variance on three language measures, but only one measure of phonological awareness (sensitivity to word-final phonemic structure) explained any significant amount of unique variance beyond that, and on only one language measure (number of bound morphemes). Age at first implant, but no other factors related to hearing loss or its treatment, explained significant amounts of variance on the language measures, as well.
In spite of early intervention and advances in implant technology, children with CIs are still delayed in learning language, but grammatical knowledge is less affected than phonological awareness. Because there was little contribution to language development measured for phonological awareness independent of lexical knowledge, it was concluded that children with CIs could benefit from intervention focused specifically on helping them learn language structures, in spite of the likely phonological deficits they experience as a consequence of having degraded inputs.
PMCID: PMC4142107  PMID: 24992492
language; children; cochlear implants
21.  Perisaccadic Remapping and Rescaling of Visual Responses in Macaque Superior Colliculus 
PLoS ONE  2012;7(12):e52195.
Visual neurons have spatial receptive fields that encode the positions of objects relative to the fovea. Because foveate animals execute frequent saccadic eye movements, this position information is constantly changing, even though the visual world is generally stationary. Interestingly, visual receptive fields in many brain regions have been found to exhibit changes in strength, size, or position around the time of each saccade, and these changes have often been suggested to be involved in the maintenance of perceptual stability. Crucial to the circuitry underlying perisaccadic changes in visual receptive fields is the superior colliculus (SC), a brainstem structure responsible for integrating visual and oculomotor signals. In this work we have studied the time-course of receptive field changes in the SC. We find that the distribution of the latencies of SC responses to stimuli placed outside the fixation receptive field is bimodal: The first mode is comprised of early responses that are temporally locked to the onset of the visual probe stimulus and stronger for probes placed closer to the classical receptive field. We suggest that such responses are therefore consistent with a perisaccadic rescaling, or enhancement, of weak visual responses within a fixed spatial receptive field. The second mode is more similar to the remapping that has been reported in the cortex, as responses are time-locked to saccade onset and stronger for stimuli placed in the postsaccadic receptive field location. We suggest that these two temporal phases of spatial updating may represent different sources of input to the SC.
PMCID: PMC3524080  PMID: 23284931
22.  Eye movement impairments in Parkinson's disease: possible role of extradopaminergic mechanisms 
BMC Neurology  2012;12:5.
The basal ganglia (BG) are thought to play an important role in the control of eye movements. Accordingly, the broad variety of subtle oculomotor alterations that has been described in Parkinson's disease (PD) are generally attributed to the dysfunction of the BG dopaminergic system. However, the present study suggest that dopamine substitution is much less effective in improving oculomotor performance than it is in restoring skeletomotor abilities.
We investigated reactive, visually guided saccades (RS), smooth pursuit eye movements (SPEM), and rapidly left-right alternating voluntary gaze shifts (AVGS) by video-oculography in 34 PD patients receiving oral dopaminergic medication (PD-DA), 14 patients with deep brain stimulation of the nucleus subthalamicus (DBS-STN), and 23 control subjects (CTL);In addition, we performed a thorough review of recent literature according therapeuthic effects on oculomotor performance in PD by switching deep brain stimulation off and on in the PD-DBS patients, we achieved swift changes between their therapeutic states without the delays of dopamine withdrawal. In addition, participants underwent neuropsychological testing.
Patients exhibited the well known deficits such as increased saccade latency, reduced SPEM gain, and reduced frequency and amplitude of AVGS. Across patients none of the investigated oculomotor parameters correlated with UPDRS III whereas there was a negative correlation between SPEM gain and susceptibility to interference (Stroop score). Of the observed deficiencies, DBS-STN slightly improved AVGS frequency but neither AVGS amplitude nor SPEM or RS performance.
We conclude that the impairment of SPEM in PD results from a cortical, conceivably non-dopaminergic dysfunction, whereas patients' difficulty to rapidly execute AVGS might be related to their BG dysfunction.
PMCID: PMC3306761  PMID: 22375860
Deep brain stimulation; Parkinson's Disease; Oculomotor function; Neurophysiology; Eye movement; Neurodegeneration
23.  Age-related decline of peripheral visual processing: the role of eye movements 
Earlier work suggests that the area of space from which useful visual information can be extracted (useful field of view, UFoV) shrinks in old age. We investigated whether this shrinkage, documented previously with a visual search task, extends to a bimanual tracking task. Young and elderly subjects executed two concurrent tracking tasks with their right and left arms. The separation between tracking displays varied from 3 to 35 cm. Subjects were asked to fixate straight ahead (condition FIX) or were free to move their eyes (condition FREE). Eye position was registered. In FREE, young subjects tracked equally well at all display separations. Elderly subjects produced higher tracking errors, and the difference between age groups increased with display separation. Eye movements were comparable across age groups. In FIX, elderly and young subjects tracked less well at large display separations. Seniors again produced higher tracking errors in FIX, but the difference between age groups did not increase reliably with display separation. However, older subjects produced a substantial number of illicit saccades, and when the effect of those saccades was factored out, the difference between young and older subjects’ tracking did increase significantly with display separation in FIX. We conclude that the age-related shrinkage of UFoV, previously documented with a visual search task, is observable with a manual tracking task as well. Older subjects seem to partly compensate their deficit by illicit saccades. Since the deficit is similar in both conditions, it may be located downstream from the convergence of retinal and oculomotor signals.
PMCID: PMC3279647  PMID: 22179529
Aging; Attention; Oculomotor control; Arm motor control; Tracking; Dual-task
24.  Relation of Frontal Eye Field Activity to Saccade Initiation during a Countermanding Task 
The countermanding (or stop signal) task probes the control of the initiation of a movement by measuring subjects’ ability to withhold a movement in various degrees of preparation in response to an infrequent stop signal. Previous research found that saccades are initiated when the activity of movement-related neurons reaches a threshold, and saccades are withheld if the growth of activity is interrupted. To extend and evaluate this relationship of frontal eye field (FEF) activity to saccade initiation, two new analyses were performed. First, we fit a neurometric function that describes the proportion of trials with a stop signal in which neural activity exceeded a criterion discharge rate as a function of stop signal delay, to the inhibition function that describes the probability of producing a saccade as a function of stop signal delay. The activity of movement-related but not visual neurons provided the best correspondence between neurometric and inhibition functions. Second, we determined the criterion discharge rate that optimally discriminated between the distributions of discharge rates measured on trials when saccades were produced or withheld. Differential activity of movement-related but not visual neurons could distinguish whether a saccade occurred. The threshold discharge rates determined for individual neurons through these two methods agreed. To investigate how reliably movement-related activity predicted movement initiation; the analyses were carried out with samples of activity from increasing numbers of trials from the same or from different neurons. The reliability of both measures of initiation threshold improved with number of trials and neurons to an asymptote of between 10 to 20 movement-related neurons. Combining the activity of visual neurons did not improve the reliability of predicting saccade initiation. These results demonstrate how the activity of a population of movement-related but not visual neurons in the FEF contributes to the control of saccade initiation. The results also validate these analytical procedures for identifying signals that control saccade initiation in other brain structures.
PMCID: PMC2748998  PMID: 18604527
25.  Role of Arcuate Frontal Cortex of Monkeys in Smooth Pursuit Eye Movements. I. Basic Response Properties to Retinal Image Motion and Position 
Journal of neurophysiology  2002;87(6):2684-2699.
Anatomical and physiological studies have shown that the “frontal pursuit area” (FPA) in the arcuate cortex of monkeys is involved in the control of smooth pursuit eye movements. To further analyze the signals carried by the FPA, we examined the activity of pursuit-related neurons recorded from a discrete region near the arcuate spur during a variety of oculomotor tasks. Pursuit neurons showed direction tuning with a wide range of preferred directions and a mean full width at half-maximum of 129°. Analysis of latency using the “receiver operating characteristic” to compare responses to target motion in opposite directions showed that the directional response of 58% of FPA neurons led the initiation of pursuit, while 19% led by 25 ms or more. Analysis of neuronal responses during pursuit of a range of target velocities revealed that the sensitivity to eye velocity was larger during the initiation of pursuit than during the maintenance of pursuit, consistent with two components of firing related to image motion and eye motion. FPA neurons showed correlates of two behavioral features of pursuit documented in prior reports. 1) Eye acceleration at the initiation of pursuit declines as a function of the eccentricity of the moving target. FPA neurons show decreased firing at the initiation of pursuit in parallel with the decline in eye acceleration. This finding is consistent with prior suggestions that the FPA plays a role in modulating the gain of visual-motor transmission for pursuit. 2) A stationary eccentric cue evokes a smooth eye movement opposite in direction to the cue and enhances the pursuit evoked by subsequent target motions. Many pursuit neurons in the FPA showed weak, phasic visual responses for stationary targets and were tuned for the positions about 4° eccentric on the side opposite to the preferred pursuit direction. However, few neurons (12%) responded during the preparation or execution of saccades. The responses to the stationary target could account for the behavioral effects of stationary, eccentric cues. Further analysis of the relationship between firing rate and retinal position error during pursuit in the preferred and opposite directions failed to provide evidence for a large contribution of image position to the firing of FPA neurons. We conclude that FPA processes information in terms of image and eye velocity and that it is functionally separate from the saccadic frontal eye fields, which processes information in terms of retinal image position.
PMCID: PMC2652416  PMID: 12037171

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