Spatial bias in anti-saccade endpoints following bilateral dorsal posterior parietal lesions.

Anti-saccades are eye movements in which the saccade is executed in the opposite direction of a visual target and are often hypometric. Because the visual target and saccade goal are decoupled, it has been suggested that competition between the two locations occurs and needs to be resolved. It has been hypothesized that the hypometria of anti-saccades reflects this spatial competition by revealing a bias towards the visual target. To confirm that this hypometria is not simply due to reduced gain, we tested 10 healthy subjects on three different anti-saccade spatial configuration tasks: 90° away across hemifields, 90° away within the same hemifield and 180° away (classic, diagonally opposite). Specifically, we examined whether saccade endpoints showed evidence for the visual target location's interference with anti-saccade programming and execution processes. Among other neural substrates involved in anti-saccades production, the dorsal posterior parietal cortex (PPC) has been implicated in the spatial inhibition of contralateral visual target. To gain insight into the neural processes involved in spatial competition during anti-saccades, we also tested one patient with a bilateral dorsal PPC lesion. In all spatial configurations, we observed that anti-saccade endpoints demonstrated a spatial bias towards the visual target for all participants, likely due to an incomplete inhibition of the visual target location. This spatial bias was exacerbated in our patient, which suggests that the dorsal PPC contributes to the amalgamation of the two competing spatial representations.

Pre-saccadic attention relies more on suppression than does covert attention.

During covert and pre-saccadic attentional shifts, it is unclear how facilitation and suppression processes interact for target selection. A recent countermanding task pointed to greater suppression at unattended locations during trials with saccades compared to trials without saccades (i.e., fixation and successful stop trials), whereas target facilitation did not differ. It is unknown whether this finding is restricted to countermanding paradigms that involve inhibitory processes. To test this, we adapted Gaspelin and colleagues (2015)'s attention capture task where, within the same block, one location was primed with frequent line discrimination trials, and all locations were occasionally probed using letters report trials. Participants also performed a baseline condition without priming. We tested 15 participants and examined how performance at non-primed locations was affected by covert versus pre-saccadic attention in blocks of four or six items, as well as by position from the primed location and timing from saccade onset. For both attention conditions, letter report at non-primed locations was worse compared to baseline, demonstrating suppression, and letter report at primed location was better, demonstrating facilitation. In saccades trials, letter report was better at primed locations and worse at non-primed locations compared to fixation trials. The timing of this additional pre-saccadic suppression differed from saccadic suppression. In both attention conditions, suppression was greater when primed and non-primed locations were within the same hemifield or in diagonal opposite quadrants. These results confirmed that attention preceding saccade execution suppressed non-primed locations to a larger extent than covert attention, with the same spatial quadrant effect.

Impaired Spatial Inhibition Processes for Interhemispheric Anti-saccades following Dorsal Posterior Parietal Lesions.

Anti-saccades are eye movements that require inhibition to stop the automatic saccade to the visual target and to perform instead a saccade in the opposite direction. The inhibitory processes underlying anti-saccades have been primarily associated with frontal cortex areas for their role in executive control. Impaired performance in anti-saccades has also been associated with the parietal cortex, but its role in inhibitory processes remains unclear. Here, we tested the assumption that the dorsal parietal cortex contributes to spatial inhibition processes of contralateral visual target. We measured anti-saccade performance in 2 unilateral optic ataxia patients and 15 age-matched controls. Participants performed 90 degree (across and within visual fields) and 180 degree inversion anti-saccades, as well as pro-saccades. The main result was that our patients took longer to inhibit visually guided saccades when the visual target was presented in the ataxic hemifield and the task required a saccade across hemifields. This was observed through anti-saccades latencies and error rates. These deficits show the crucial role of the dorsal posterior parietal cortex in spatial inhibition of contralateral visual target representations to plan an accurate anti-saccade toward the ipsilesional side.

Movement drift in optic ataxia reveals deficits in hand state estimation in oculocentric coordinates.

When vision is removed, limb position has been shown to progressively drift during repetitive arm movements. The posterior parietal cortex (PPC) is known to be involved in the processing of multisensory information, the formation of internal hand estimate, and online motor control. Here, we compared hand position drift between healthy controls and 2 patients with PPC damage to gain insight into the mechanisms underlying movement drift and investigate the possible role of the PPC in this process. To do so, we asked participants to perform back-and-forth movements between 2 targets, in the dark and under different gaze fixation conditions. Each individual participant consistently drifted to the same end position for a given hand and gaze condition. We found that the final drift distance was related to small systematic errors made on the very first trial in the dark, with an approximate 3.5 fold increase in magnitude. Furthermore, PPC damage resulted in greater movement drift in patients when the unseen hand was in the contralesional oculocentric space and also when the target was located in the lower visual field. We conclude that the PPC is involved in the proprioceptive representation of hand position in oculocentric coordinates used for reach planning and motor control. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Predicted Position Error Triggers Catch-Up Saccades during Sustained Smooth Pursuit.

For humans, visual tracking of moving stimuli often triggers catch-up saccades during smooth pursuit. The switch between these continuous and discrete eye movements is a trade-off between tolerating sustained position error (PE) when no saccade is triggered or a transient loss of vision during the saccade due to saccadic suppression. de Brouwer et al. (2002b) demonstrated that catch-up saccades were less likely to occur when the target re-crosses the fovea within 40-180 ms. To date, there is no mechanistic explanation for how the trigger decision is made by the brain. Recently, we proposed a stochastic decision model for saccade triggering during visual tracking (Coutinho et al., 2018) that relies on a probabilistic estimate of predicted PE (PEpred). Informed by model predictions, we hypothesized that saccade trigger time length and variability will increase when pre-saccadic predicted errors are small or visual uncertainty is high (e.g., for blurred targets). Data collected from human participants performing a double step-ramp task showed that large pre-saccadic PEpred (>10°) produced short saccade trigger times regardless of the level of uncertainty while saccade trigger times preceded by small PEpred (<10°) significantly increased in length and variability, and more so for blurred targets. Our model also predicted increased signal-dependent noise (SDN) as retinal slip (RS) increases; in our data, this resulted in longer saccade trigger times and more smooth trials without saccades. In summary, our data supports our hypothesized predicted error-based decision process for coordinating saccades during smooth pursuit.

Spatial and temporal dynamics of presaccadic attentional facilitation before pro- and antisaccades.

The premotor theory of attention and the visual attention model make different predictions about the temporal and spatial allocation of presaccadic attentional facilitation. The current experiment investigated the spatial and temporal dynamics of presaccadic attentional facilitation during pro- and antisaccade planning; we investigated whether attention shifts only to the saccade goal location or to the target location or elsewhere, and when. Participants performed a dual-task paradigm with blocks of either anti- or prosaccades and also discriminated symbols appearing at different locations before saccade onset (measure of attentional allocation). In prosaccades blocks, correct prosaccade discrimination was best at the target location, while during errors, discrimination was best at the location opposite to the target location. This pattern was inversed in antisaccades blocks, although discrimination remained high opposite to the target location. In addition, we took the benefit of a large range of saccadic landing positions and showed that performance across both types of saccades was best at the actual saccade goal location (where the eye will actually land) rather than at the instructed position. Finally, temporal analyses showed that discrimination remained highest at the saccade goal location, from long before to closer to saccade onset, increasing slightly for antisaccades closer to saccade onset. These findings are in line with the premises of the premotor theory of attention, showing that attentional allocation is primarily linked both temporally and spatially to the saccade goal location.

Contrasting effects of exogenous cueing on saccades and reaches.

Previous studies have shown that eye and arm movements tend to be intrinsically coupled in their behavior. There is, however, no consensus on whether planning of eye and arm movements is based on shared or independent representations. One way to gain insight into these processes is to compare how exogenous attentional modulation influences the temporal and spatial characteristics of the eye and the arm during single or combined movements. Thirteen participants (M = 22.8 years old, SD = 1.5) performed single or combined movements to an eccentric target. A behaviorally irrelevant cue flashed just before the target at different locations. There was no effect of the cue on the saccade or reach amplitudes, whether they were performed alone or together. We found no differences in overall reaction times (RTs) between single and combined movements. With respect to the effect of the cue, both saccades and reaches followed a similar pattern with the shortest RTs when the cue was closest to the target, which we propose reflects effector-independent processes. Compared to when no cue was presented before the target, saccade RTs were generally inhibited by the irrelevant cue with increasing cue-target distance. In contrast, reach RTs showed strong facilitation at the target location and less facilitation at farther distances. We propose that this reflects the presence of effector-dependent processes. The similarities and differences in RTs between the saccades and reaches are consistent with effector-dependent and -independent processes working in parallel.

Learned rather than online relative weighting of visual-proprioceptive sensory cues.

When reaching to an object, information about the target location as well as the initial hand position is required to program the motor plan for the arm. The initial hand position can be determined by proprioceptive information as well as visual information, if available. Bayes-optimal integration posits that we utilize all information available, with greater weighting on the sense that is more reliable, thus generally weighting visual information more than the usually less reliable proprioceptive information. The criterion by which information is weighted has not been explicitly investigated; it has been assumed that the weights are based on task- and effector-dependent sensory reliability requiring an explicit neuronal representation of variability. However, the weights could also be determined implicitly through learned modality-specific integration weights and not on effector-dependent reliability. While the former hypothesis predicts different proprioceptive weights for left and right hands, e.g., due to different reliabilities of dominant vs. nondominant hand proprioception, we would expect the same integration weights if the latter hypothesis was true. We found that the proprioceptive weights for the left and right hands were extremely consistent regardless of differences in sensory variability for the two hands as measured in two separate complementary tasks. Thus we propose that proprioceptive weights during reaching are learned across both hands, with high interindividual range but independent of each hand's specific proprioceptive variability. NEW & NOTEWORTHY How visual and proprioceptive information about the hand are integrated to plan a reaching movement is still debated. The goal of this study was to clarify how the weights assigned to vision and proprioception during multisensory integration are determined. We found evidence that the integration weights are modality specific rather than based on the sensory reliabilities of the effectors.

Corrective response times in a coordinated eye-head-arm countermanding task.

Inhibition of motor responses has been described as a race between two competing decision processes of motor initiation and inhibition, which manifest as the reaction time (RT) and the stop signal reaction time (SSRT); in the case where motor initiation wins out over inhibition, an erroneous movement occurs that usually needs to be corrected, leading to corrective response times (CRTs). Here we used a combined eye-head-arm movement countermanding task to investigate the mechanisms governing multiple effector coordination and the timing of corrective responses. We found a high degree of correlation between effector response times for RT, SSRT, and CRT, suggesting that decision processes are strongly dependent across effectors. To gain further insight into the mechanisms underlying CRTs, we tested multiple models to describe the distribution of RTs, SSRTs, and CRTs. The best-ranked model (according to 3 information criteria) extends the LATER race model governing RTs and SSRTs, whereby a second motor initiation process triggers the corrective response (CRT) only after the inhibition process completes in an expedited fashion. Our model suggests that the neural processing underpinning a failed decision has a residual effect on subsequent actions. NEW & NOTEWORTHY Failure to inhibit erroneous movements typically results in corrective movements. For coordinated eye-head-hand movements we show that corrective movements are only initiated after the erroneous movement cancellation signal has reached a decision threshold in an accelerated fashion.

Transsaccadic memory of multiple spatially variant and invariant object features.

Transsaccadic memory is a process by which remembered object information is updated across a saccade. To date, studies on transsaccadic memory have used simple stimuli-that is, a single dot or feature of an object. It remains unknown how transsaccadic memory occurs for more realistic, complex objects with multiple features. An object's location is a central feature for transsaccadic updating, as it is spatially variant, but other features such as size are spatially invariant. How these spatially variant and invariant features of an object are remembered and updated across saccades is not well understood. Here we tested how well 14 participants remembered either three different features together (location, orientation, and size) or a single feature at a time of a bar either while fixating either with or without an intervening saccade. We found that the intervening saccade influenced memory of all three features, with consistent biases of the remembered location, orientation, and size that were dependent on the direction of the saccade. These biases were similar whether participants remembered either a single feature or multiple features and were not observed with increased memory load (single vs. multiple features during fixation trials), confirming that these effects were specific to the saccade updating mechanisms. We conclude that multiple features of an object are updated together across eye movements, supporting the notion that spatially invariant features of an object are bound to their location in memory.

Effects of a pretarget distractor on saccade reaction times across space and time in monkeys and humans.

Previous studies have shown that the influence of a behaviorally irrelevant distractor on saccade reaction times (SRTs) varies depending on the temporal and spatial relationship between the distractor and the saccade target. We measured distractor influence on SRTs to a subsequently presented target, varying the spatial location and the timing between the distractor and the target. The distractor appeared at one of four equally eccentric locations, followed by a target (either 50 ms or 200 ms after) at one of 136 different locations encompassing an area of 20° square. We extensively tested two humans and two monkeys on this task to determine interspecies similarities and differences, since monkey neurophysiology is often used to interpret human behavioral findings. Results were similar across species; for the short interval (50 ms), SRTs were shortest to a target presented close to or at the distractor location and increased primarily as a function of the distance from the distractor. There was also an effect of distractor-target direction and visual field. For the long interval (200 ms) the results were inverted; SRTs were longest for short distances between the distractor and target and decreased as a function of distance from distractor. Both SRT patterns were well captured by a two-dimensional dynamic field model with short-distance excitation and long-distance inhibition, based upon known functional connectivity found in the superior colliculus that includes wide-spread excitation and inhibition. Based on these findings, we posit that the different time-dependent patterns of distractor-related SRTs can emerge from the same underlying neuronal mechanisms common to both species.

Saccade execution suppresses discrimination at distractor locations rather than enhancing the saccade goal location.

As we have limited processing abilities with respect to the plethora of visual information entering our brain, spatial selection mechanisms are crucial. These mechanisms result in both enhancing processing at a location of interest and in suppressing processing at other locations; together, they enable successful further processing of locations of interest. It has been suggested that saccade planning modulates these spatial selection mechanisms; however, the precise influence of saccades on the distribution of spatial resources underlying selection remains unclear. To this end, we compared discrimination performance at different locations (six) within a work space during different saccade tasks. We used visual discrimination performance as a behavioral measure of enhancement and suppression at the different locations. A total of 14 participants performed a dual discrimination/saccade countermanding task, which allowed us to specifically isolate the consequences of saccade execution. When a saccade was executed, discrimination performance at the cued location was never better than when fixation was maintained, suggesting that saccade execution did not enhance processing at a location more than knowing the likelihood of its appearance. However, discrimination was consistently lower at distractor (uncued) locations in all cases where a saccade was executed compared with when fixation was maintained. Based on these results, we suggest that saccade execution specifically suppresses distractor locations, whereas attention shifts (with or without an accompanying saccade) are involved in enhancing perceptual processing at the goal location.

Motion integration for ocular pursuit does not hinder perceptual segregation of moving objects.

When confronted with a complex moving stimulus, the brain can integrate local element velocities to obtain a single motion signal, or segregate the elements to maintain awareness of their identities. The integrated motion signal can drive smooth-pursuit eye movements (Heinen and Watamaniuk, 1998), whereas the segregated signal guides attentive tracking of individual elements in multiple-object tracking tasks (MOT; Pylyshyn and Storm, 1988). It is evident that these processes can occur simultaneously, because we can effortlessly pursue ambulating creatures while inspecting disjoint moving features, such as arms and legs, but the underlying mechanism is unknown. Here, we provide evidence that separate neural circuits perform the mathematically opposed operations of integration and segregation, by demonstrating with a dual-task paradigm that the two processes do not share attentional resources. Human observers attentively tracked a subset of target elements composing a small MOT stimulus, while pursuing it ocularly as it translated across a computer display. Integration of the multidot stimulus yielded optimal pursuit. Importantly, performing MOT while pursuing the stimulus did not degrade performance on either task compared with when each was performed alone, indicating that they did not share attention. A control experiment showed that pursuit was not driven by integration of only the nontargets, leaving the MOT targets free for segregation. Nor was a predictive strategy used to pursue the stimulus, because sudden changes in its global velocity were accurately followed. The results suggest that separate neural mechanisms can simultaneously segregate and integrate the same motion signals.

Testing for optic ataxia in a blind field.

Optic ataxia is a component of Balint's syndrome and is a disorder that results from damage to the posterior parietal cortex (PPC) leading to deficits in reaching and grasping objects presented in the visual field opposite to the damaged hemisphere. It is also often the case that Balint's syndrome is accompanied by visual field defects due to the proximity of parietal and occipital cortices and also due to the subcortical pathway relaying visual information from the retina to the visual cortex passing underneath the parietal cortex. The presence of primary visual defects such as hemianopia often prevents clinicians from diagnosing higher-level visual deficits such as optic ataxia; the patient cannot reach to targets he/she cannot see. Here, we show that through the use of a paradigm that takes advantage of remapping mechanisms, we were able to observe optic ataxia in the blind field. We measured reach endpoints of a patient presenting with left optic ataxia as well as a quadrantanopia in the left lower visual field in eye-static and eye-dynamic conditions. In static conditions, we first asked the patient to reach to targets viewed in her non-optic ataxic intact right visual field (fixating on the left of the target array). In this case, the patient showed undershoots equivalent to controls. Next, we asked her to reach to (the same) targets viewed in the upper left optic ataxic but intact visual field (fixating to the right of the target array). The undershooting pattern increased greatly, consistent with unilateral left optic ataxia. In dynamic conditions, we asked her to view targets in her good (right lower) visual field before reorienting her line of sight to the opposite side, causing the internal representation of the target to be updated into the opposite (ataxic) blind visual field. The patient then reached to the remembered (and updated) location of the target. We found errors typical of optic ataxia for reaches guided toward the quadrantanopic field. This confirmed that reaching errors depended on the updated internal representation of the target and not on where the target was viewed initially. In both the patient and the controls, the updating of target location was partial, with reaching errors observed subsequent to an eye movement made from left to right fixation positions being intermediate between the left and right static conditions. Thus, using this remapping paradigm, we were able to observe optic ataxia in the blind field. In conclusion, this remapping paradigm would allow clinicians to test for visuo-manual transformation deficits (optic ataxia) even when it is associated with hemianopia.

Causal evidence for posterior parietal cortex involvement in visual-to-motor transformations of reach targets.

It has been posited that the posterior parietal cortex (PPC) is involved in the visual-to-motor transformation for reach planning. Such a transformation is required because in general the retinal information and the arm motor command do not align, for example in the case of non-zero eye/head orientations. Here, we present behavioral data from a patient with unilateral optic ataxia consecutive to damage to the superior parietal lobule including the intraparietal sulcus in the right hemisphere, who we asked to reach to visual targets under different head roll angles. An accurate visual-to-motor transformation has to integrate head roll to compensate for the rotated retinal location of the target, resulting in a head roll-independent pattern of reach endpoints. If however, head roll is not compensated for, reach endpoints should vary across different head rolls, reflecting a reach plan based on the rotated retinal target location. Remarkably, the patient compensated for head roll when reaching to targets presented within his intact right visual field (VF) (not different from controls) but not for reaches to targets in the contralesional left VF. The amount of compensation was the same irrespective of whether the initial hand position was located in the left or right VF, showing that this transformation concerns only the target location and not the hand-target motor vector. We interpret these findings as causal evidence for the involvement of the PPC in integrating head roll signals in the visual-to-motor transformation of the reach target.

The eye dominates in guiding attention during simultaneous eye and hand movements.

Prior to the onset of a saccade or a reach, attention is directed to the goal of the upcoming movement. However, it remains unknown whether attentional resources are shared across effectors for simultaneous eye and hand movements. Using a 4-AFC shape discrimination task, we investigated attentional allocation during the planning of a saccade alone, reach alone, or combined saccade and reach to one of five peripheral locations. Target discrimination was better when the probe appeared at the goal of the impending movement than when it appeared elsewhere. However, discrimination performance at the movement goal was not better for combined eye-hand movements compared to either effector alone, suggesting a shared limited attentional resource rather than separate pools of effector-specific attention. To test which effector dominates in guiding attention, we then separated eye and hand movement goals in two conditions: (1) cued reach/fixed saccade--subjects made saccades to the same peripheral location throughout the block, while the reach goal was cued and (2) cued saccade/fixed reach--subjects made reaches to the same location, while the saccade goal was cued. For both conditions, discrimination performance was consistently better at the eye goal than the hand goal. This indicates that shared attentional resources are guided predominantly by the eye during the planning of eye and hand movements.

The default allocation of attention is broadly ahead of smooth pursuit.

When moving through our environment, it is vital to preferentially process positions on our future path in order to react quickly to critical situations. During smooth pursuit, attention may be directed ahead with either a focused locus or a broad bias. We examined the 2D spatial extent of attention during a smooth pursuit task using both saccade (SRT) and manual (MRT) reaction times as measures of attentional allocation. Targets were flashed at various locations around current eye position while subjects pursued a moving target. Subjects made a saccade or pressed a button as soon as they perceived the target. Both SRTs and MRTs were shortest to targets flashed ahead of compared to behind the direction of pursuit across half of the visual field ahead of pursuit direction. Furthermore, we found an increase specific to SRTs at small target eccentricities directly ahead of pursuit, which may be related to an additional saccade trigger strategy; small saccades take longer to execute if smooth pursuit brings the eyes close to the target. In summary, both SRTs and MRTs revealed that attention is by default broadly allocated in the visual hemi-field ahead of the line of sight during smooth pursuit eye movements. This attentional bias may serve a predictive purpose for facilitating the processing of upcoming events.

Attentional cueing at the saccade goal, not at the target location, facilitates saccades.

Presenting a behaviorally irrelevant cue shortly before a target at the same location decreases the latencies of saccades to the target, a phenomenon known as exogenous attention facilitation. It remains unclear whether exogenous attention interacts with early, sensory stages or later, motor planning stages of saccade production. To distinguish between these alternatives, we used a saccadic adaptation paradigm to dissociate the location of the visual target from the saccade goal. Three male and four female human subjects performed both control trials, in which saccades were made to one of two target eccentricities, and adaptation trials, in which the target was shifted from one location to the other during the saccade. This manipulation adapted saccades so that they eventually were directed to the shifted location. In both conditions, a behaviorally irrelevant cue was flashed 66.7 ms before target appearance at a randomly selected one of seven positions that included the two target locations. In control trials, saccade latencies were shortest when the cue was presented at the target location and increased with cue-target distance. In contrast, adapted saccade latencies were shortest when the cue was presented at the adapted saccade goal, and not at the visual target location. The dynamics of adapted saccades were also altered, consistent with prior adaptation studies, except when the cue was flashed at the saccade goal. Overall, the results suggest that attentional cueing facilitates saccade planning rather than visual processing of the target.

Pre-saccadic perceptual facilitation can occur without covert orienting of attention.

The pre-motor theory of attention suggests that the mechanisms involved in target selection for eye movements are the same as those for spatial attention shifts. The pre-saccadic facilitation of perceptual discrimination at the location of a saccadic goal (paradigm of Deubel and Schneider, 1996) has been considered as an argument for this theory. We compared letter discrimination performance in a saccade (overt attention - pre-saccadic facilitation) and a fixation (covert attention) task in a patient with right posterior parietal damage and 4 controls. In the overt attention condition, the patient was instructed by a central cue to make a saccade to a target located at a peripheral location. During the saccade latency (in a period of time of 250 msec following the presentation of the cue), a letter was presented at the target location. Accuracy of leftward saccades was impaired compared to rightward saccades. To evaluate letter discrimination performance in this saccade task (i.e., the presence of pre-saccadic facilitation), we selected only those leftward saccades that were equivalent in accuracy (and latency) to the rightward ones. Within these selected trials, the patient was able to discriminate letters equally well in both visual fields. In contrast, he performed at chance level during the fixation task (covert attention condition) for letters presented at the same peripheral location with the same timing with respect to the cue presentation. The patient could thus discriminate the letter presented at 8° of visual eccentricity while he was preparing a saccade, whereas he was unable to perceive the letter in the fixation task. Remarkably, in the left visual field, letter discrimination was impossible even when a letter was presented as close as 2.5° of visual eccentricity in the fixation task. Altogether, these results suggest that pre-saccadic perceptual facilitation does not rely on the same processes as those of covert attention, as tested by fixation task. Instead, we propose that pre-saccadic perceptual facilitation results from a form of attention specific to action, which could correspond to a pre-saccadic remapping process.

Differential influence of attention on gaze and head movements.

A salient peripheral cue can capture attention, influencing subsequent responses to a target. Attentional cueing effects have been studied for head-restrained saccades; however, under natural conditions, the head contributes to gaze shifts. We asked whether attention influences head movements in combined eye-head gaze shifts and, if so, whether this influence is different for the eye and head components. Subjects made combined eye-head gaze shifts to horizontal visual targets. Prior to target onset, a behaviorally irrelevant cue was flashed at the same (congruent) or opposite (incongruent) location at various stimulus-onset asynchrony (SOA) times. We measured eye and head movements and neck muscle electromyographic signals. Reaction times for the eye and head were highly correlated; both showed significantly shorter latencies (attentional facilitation) for congruent compared with incongruent cues at the two shortest SOAs and the opposite pattern (inhibition of return) at the longer SOAs, consistent with attentional modulation of a common eye-head gaze drive. Interestingly, we also found that the head latency relative to saccade onset was significantly shorter for congruent than that for incongruent cues. This suggests an effect of attention on the head separate from that on the eyes.