Microsaccades keep the eyes' balance during fixation

During fixation of a stationary target, small involuntary eye movements exhibit an erratic trajectory-a random walk. Two types of these fixational eye movements are drift and microsaccades (small-amplitude saccades). We investigated fixational eye movements and binocular coordination using a statistical analysis that had previously been applied to human posture control. This random-walk analysis uncovered two different time scales in fixational eye movements and identified specific functions for microsaccades. On a short time scale, microsaccades enhanced perception by increasing fixation errors. On a long time scale, microsaccades reduced fixation errors and binocular disparity (relative to pure drift movements). Thus, our findings clarify the role of oculomotor processes during fixation.     

The time course of attentional and oculomotor capture reveals a common cause

Eye movements are often misdirected toward a distractor when it appears abruptly, an effect known as oculomotor capture. Fundamental differences between eye movements and attention have led to questions about the relationship of oculomotor capture to the more general effect of sudden onsets on performance, known as attentional capture. This study explores that issue by examining the time course of eye movements and manual localization responses to targets in the presence of sudden-onset distractors. The results demonstrate that for both response types, the proportion of trials on which responses are erroneously directed to sudden onsets reflects the quality of information about the visual display at a given point in time. Oculomotor capture appears to be a specific instance of a more general attentional capture effect. Differences and similarities between the two types of capture can be explained by the critical idea that the quality of information about a visual display changes over time and that different response systems tend to access this information at different moments in time.     

The relationship between covert and overt attention in endogenous cuing

In a standard Posner paradigm, participants were endogenously cued to attend to a peripheral location in visual space without making eye movements. They responded faster to target letters presented at cued than at uncued locations. On some trials, instead of a manual response, they had to move their eyes to a location in space. Results showed that the eyes deviated away from the validly cued location; when the cue was invalid and attention had to be allocated to the uncued location, eye movements also deviated away, but now from the uncued location. The extent to which the eyes deviated from cued and uncued locations was related to the dynamics of attention allocation. We hypothesized that this deviation was due to the successful inhibition of the attended location. The results imply that the oculomotor system is not only involved during the endogenous direction of covert attention to a cued location, but also when covert attention is directed to an uncued location. It appears that the oculomotor system is activated wherever spatial attention is allocated. The strength of saccade deviation might turn out to be an important measure for the amount of attention allocated to any particular location over time.     

Attentional localization prior to simple and directed manual responses

The relationship between attention and the programming of motor responses was investigated, using a paradigm in which the onsets of targets for movements were preceded by peripheral attentional cues. Simple (button release) and reaching manual responses were compared under conditions in which the subjects either made saccades toward the target location or refrained from making eye movements. The timing of the movement onset was used as the dependent measure for both simple and reaching manual responses. Eye movement latencies were also measured. A follow-up experiment measured the effect of the same peripheral cuing procedure on purely visual processes, using signal detection measures of visual sensitivity and response bias. The results of the first experiment showed that reaction time (RT) increased with the distance between the cued and the target locations. Stronger distance effects were observed when goal-directed responses were required, which suggests enhanced attentional localization of target positions under these conditions. The requirement to generate an eye movement response was found to delay simple manual RTs. However, mean reaching RTs were unaffected by the eye movement condition. Distance gradients on eye movement latencies were relatively shallow, as compared with those on goal-directed manual responses. The second experiment showed that the peripheral cue had only a very small effect on visual detection sensitivity in the absence of directed motor responses. It is concluded that cue-target distance effects with peripheral cues are modulated by the motor-programming requirements of the task. The effect of the peripheral cue on eye movement latencies was qualitatively different from that observed on manual RTs, indicating the existence of separate neural representations underlying both response types. At the same time, the interactions between response modalities are consistent with a supramodal representation of attentional space, within which different motor programs may interact.     

Attentional localization prior to simpleand directed manual responses

The relationship between attention and the programming of motor responses was investigated, using a paradigm in which the onsets of targets for movements were preceded by peripheral attentional cues. Simple (button release) and reaching manual responses were compared under conditions in which the subjects either made saccades toward the target location or refrained from making eye movements. The timing of the movement onset was used as the dependent measure for both simple and reaching manual responses. Eye movement latencies were also measured. A follow-up experiment measured the effect of the same peripheral cuing procedure on purely visual processes, using signal detection mea-sures of visual sensitivity and response bias. The results of the first experiment showed that reaction time (RT) increased with the distance between the cued and the target locations. Stronger distance ef-fects were observed when goal-directed responses were required, which suggests enhanced attentional localization of target positions under these conditions. The requirement to generate an eye movement response was found to delay simple manual RTs. However, mean reaching RTs were unaffected by the eye movement condition. Distance gradients on eye movement latencies were relatively shallow, as compared with those on goal-directed manual responses. The second experiment showed that the peripheral cue had only a very small effect on visual detection sensitivity in the absence of directed motor responses. It is concluded that cue-target distance effects with peripheral cues are modulated by the motor-programming requirements of the task. The effect of the peripheral cue on eye movement latencies was qualitatively different from that observed on manual RTs, indicating the existence of separate neural representations underlying both response types. At the same time, the interactions be-tween response modalities are consistent with a supramodal representation of attentional space, within which different motor programs may interact.     

Variable coordination of eye and head movements during the early development of attention: A longitudinal study of infants aged 12–36 months

This longitudinal study investigated the effects of attentional development on peripheral stimulus localization by analyzing the eye and head movements of toddlers as they matured from 12 to 36 months. On each trial of an experiment, a central fixation point and a 30° peripheral stimulus were presented, such that in the gap condition the fixation disappeared 300 ms before the peripheral stimulus, whereas in the no-overlap condition it disappeared simultaneously as the peripheral stimulus, and in the overlap condition the fixation remained present when the peripheral target occurred. Results showed that eye and head movement latencies were highly correlated in all conditions and ages. However, at 12 months, head movements were as fast as eye movements, whereas during the subsequent development, eye movements became increasingly faster than head movements. These findings are indicative of a transition between 12 and 36 months due either to a change in attentional control, or to changes in the size of the visual field in which only eye movements occur.     

Why are you looking at me? It’s because I’m talking,but mostly because I’m staring or not doing much

Our attention is particularly driven toward faces, especially the eyes, and there is much debate over the factors that modulate this social attentional orienting. Most of the previous research has presented faces in isolation, and we tried to address this shortcoming by measuring people’s eye movements whilst they observe more naturalistic and varied social interactions. Participants’ eye movements were monitored whilst they watched three different types of social interactions (monologue, manual activity, active attentional misdirection), which were either accompanied by the corresponding audio as speech or by silence. Our results showed that (1) participants spent more time looking at the face when the person was giving a monologue, than when he/she was carrying out manual activities, and in the latter case they spent more time fixating on the person’s hands. (2) Hearing speech significantly increases the amount of time participants spent looking at the face (this effect was relatively small), although this was not accounted for by any increase in mouth-oriented gaze. (3) Participants spent significantly more time fixating on the face when direct eye contact was established, and this drive to establish eye contact was significantly stronger in the manual activities than during the monologue. These results highlight people’s strategic top-down control over when they attend to faces and the eyes, and support the view that we use our eyes to signal non-verbal information.     

Perceived localizations and eye movements with action‐generated and computer‐generated vanishing points of moving stimuli

When observers localize the vanishing point of a moving target, localizations are reliably displaced beyond the final position, in the direction the stimulus was travelling just prior to its offset. We examined modulations of this phenomenon through eye movements and action control over the vanishing point. In Experiment 1 with pursuit eye movements, localization errors were in movement direction, but less pronounced when the vanishing point was self‐determined by a key press of the observer. In contrast, in Experiment 2 with fixation instruction, localization errors were opposite movement direction and independent from action control. This pattern of results points at the role of eye movements, which were gathered in Experiment 3. That experiment showed that the eyes lagged behind the target at the point in time, when it vanished from the screen, but that the eyes continued to drift on the targets' virtual trajectory. It is suggested that the perceived target position resulted from the spatial lag of the eyes and of the persisting retinal image during the drift.     

Top-down influences make saccades deviate away: the case of endogenous cues

We tested a recent hypothesis suggesting that the eye deviates away from a location when top-down preparation can influence target selection. Participants had to make an eye movement to a peripheral target. Before the upcoming target, a central cue indicated the likely target location. Results show that when the target was presented at a location different from that indicated by the cue, eye movements to the target deviated away from the cued location. Because central cues are under top-down control, the present results are in line with a determining role of top-down preparation on saccade direction. These results contrast with the findings reported in a similar paradigm executed with hand movements, in which the movements were mostly initiated in the direction of the cued location. Therefore, we conclude that inhibitory effects typically observed when executing eye movements may not be observed when executing hand movements in similar conditions.     

Watch out! Directional threat-related postures cue attention and the eyes

Previous work indicates that threatening facial expressions with averted eye gaze can act as a signal of imminent danger, enhancing attentional orienting in the gazed-at direction. However, this threat-related gaze-cueing effect is only present in individuals reporting high levels of anxiety. The present study used eye tracking to investigate whether additional directional social cues, such as averted angry and fearful human body postures, not only cue attention, but also the eyes. The data show that although body direction did not predict target location, anxious individuals made faster eye movements when fearful or angry postures were facing towards (congruent condition) rather than away (incongruent condition) from peripheral targets. Our results provide evidence for attentional cueing in response to threat-related directional body postures in those with anxiety. This suggests that for such individuals, attention is guided by threatening social stimuli in ways that can influence and bias eye movement behaviour.     

Independent allocation of attention to eye and hand targets in coordinated eye-hand movements

When reaching for objects, people frequently look where they reach. This raises the question of whether the targets for the eye and hand in concurrent eye and hand movements are selected by a unitary attentional system or by independent mechanisms. We used the deployment of visual attention as an index of the selection of movement targets and asked observers to reach and look to either the same location or separate locations. Results show that during the preparation of coordinated movements, attention is allocated in parallel to the targets of a saccade and a reaching movement. Attentional allocations for the two movements interact synergistically when both are directed to a common goal. Delaying the eye movement delays the attentional shift to the saccade target while leaving attentional deployment to the reach target unaffected. Our findings demonstrate that attentional resources are allocated independently to the targets of eye and hand movements and suggest that the goals for these effectors are selected by separate attentional mechanisms.     

Visual stability with goal-directed eye and arm movements toward a target displaced during saccadic suppression

This experiment tested whether the perceived stability of the environment is altered when there is a combination of eye and visually open-loop hand movements toward a target displaced during the eye movements, i.e., during saccadic suppression. Visual-target eccentricity randomly decreased or increased during eye movements and subjects reported whether they perceived a target displacement or not, and if so, the direction of the displacement. Three experimental conditions, involving different combinations of eye and arm movements, were tested: (a) eye movements only; (b) simultaneous eye and rapid arm movements toward the target; and (c) simultaneous eye and arm movements with a restraint blocking the arm as soon as the hand left the starting position. The perceptual threshold of target displacements resulting in an increased target eccentricity was greater when subjects combined eye and arm movements toward the target object, specially for the no-restraint condition. Subjects corrected most of their arm trajectory toward the displaced target despite the short movement times (average MT = 189 ms). After the movements, the null error feedback of the hand's final position presumably overlapped the retino-oculomotor signal error and could be responsible for the deficient perception of target displacements. Thus, subjects interpreted the terminal hand positions as being within the range of the endpoint variability associated with the production of rapid arm movements rather than as a change of the environment. These results suggest that a natural strategy adopted for processing spatial information, especially in a competing situation, could favour a constancy tendency, avoiding systematic perception of a change of environment for any noise or variability at the central or peripheral levels.     

Eye movements reveal mechanisms underlying attentional biases towards threat

Mechanisms underlying attentional biases towards threat (ABTs), such as attentional avoidance and difficulty of disengagement, are still unclear. To address this issue, we recorded participants' eye movements during a dot detection task in which threatening or neutral stimuli served as peripheral cues. We evaluated response times (RTs) in trials where participants looked at the central fixation cross (not at the cues), as they were required, and number and duration of (unwanted) fixations towards threatening or neutral cues; in all analyses trait anxiety was treated as a covariate. Difficulty in attentional disengagement (longer RTs) was found when peripheral threatening stimuli were presented for 100?ms. Moreover, we observed significantly shorter (unwanted) fixations on threatening than on neutral peripheral stimuli, compatible with an avoidance bias, for longer presentation times. These findings demonstrate that, independent of trait anxiety levels, disengagement bias occurs without eye movements, whereas eye movements are implied in threat avoidance.     

Differential use of vocal and gestural communication by chimpanzees (Pan troglodytes) in response to the attentional status of a human (Homo sapiens)

This study examined the communicative behavior of 49 captive chimpanzees (Pan troglodytes), particularly their use of vocalizations, manual gestures, and other auditory- or tactile-based behaviors as a means of gaining an inattentive audience's attention. A human (Homo sapiens) experimenter held a banana while oriented either toward or away from the chimpanzee. The chimpanzees' behavior was recorded for 60 s. Chimpanzees emitted vocalizations faster and were more likely to produce vocalizations as their 1st communicative behavior when a human was oriented away from them. Chimpanzees used manual gestures more frequently and faster when the human was oriented toward them. These results replicate the findings of earlier studies on chimpanzee gestural communication and provide new information about the intentional and functional use of their vocalizations.     

Influence of attentional capture on oculomotor control

Previous research has shown that when searching for a color singleton, top-down control cannot prevent attentional capture by an abrupt visual onset. The present research addressed whether a task-irrelevant abrupt onset would affect eye movement behavior when searching for a color singleton. Results show that in many instances the eye moved in the direction of the task-irrelevant abrupt onset. There was evidence that top-down control could neither entirely prevent attentional capture by visual onsets nor prevent the eye from starting to move in the direction of the onset. Results suggest parallel programming of 2 saccades: 1 voluntary goal-directed eye movement toward the color singleton target and 1 stimulus-driven eye movement reflexively elicited by the abrupt onset. A neurophysiologically plausible model that can account for the current findings is discussed.     

The relationship between eye movements and spatial attention

Most previous studies of the attentional consequences of making saccadic eye movements have used peripheral stimuli to elicit eye movements. It is argued that in the light of evidence showing automatic “capture” of attention by peripheral stimuli, these experiments do not distinguish between attentional effects due to peripheral stimuli and those due to eye movements. In the present study, spatial attention was manipulated by varying the probability that peripheral probe stimuli would appear in different positions, while saccades were directed by a central arrow, enabling the effects of attention and eye movements to be separated. The results showed that the time to react to a peripheral stimulus could be shortened both by advance knowledge of its likely position and, separately, by preparing to make a saccade to that position. When the saccade was directed away from the most likely position of the probe, the targets for attention and eye movements were on opposite sides of the display. In this condition, the effects of preparing to make a saccade proved to be stronger than the effects of attentional allocation until well after the saccade had finished, suggesting that making a saccade necessarily involves the allocation of attention to the target position. The effects of probe stimuli on saccade latencies were also examined: probe stimuli that appeared before the saccade shortened saccade latencies if they appeared at the saccade target, and lengthened saccade latencies if they appeared on the opposite side of fixation. These facilitatory and inhibitory effects were shown to occur at different stages of saccade preparation and suggest that attention plays an important role in the generation of voluntary eye movements. The results of this study indicate that while it is possible to make attention movements without making corresponding eye movements, it is not possible to make an eye movement (in the absence of peripheral stimulation) without making a corresponding shift in the focus of attention.     

Saccadic localization of visual targets by the very young human infant

The direction, latency, and form of the 1- and 2-month-old human infant’s saccadic eye movements toward peripheral targets were investigated. Infants of both ages reliably executed a directionally appropriate first saccade toward a peripheral target introduced as far as 30 deg from the line of sight along the horizontal and both diagonal axes, but only to 10 deg along the vertical axis. The presence of a second target in the central visual field reduced the probability of peripheral target localization. A significant inverse relation was found between target distance from the line of sight and probability of initiating a directionally appropriate saccade. Electro-oculography revealed that latency to first saccade, although highly variable, was less than 500 msec on a significant proportion of trials. Unlike the adult, the first saccade to target was grossly hypometric and was followed by one or more saccades of approximately equal amplitude to the first.     

An investigation of the relationship between eye and retinal image movement in the perception of movement

The question investigated was whether or not eye movements accompanied by abnormal retinal image movements, movements that are either or both at a different rate or in a different direction than the eye movement, predictably lead to perceived movement. Os reported whether or not they saw a visual target move when the movement of the target was either dependent on and simultaneous with their eye movements or when the target movement was independent of their eye movements. In the main experiment, observations were made when the ratio between eye and target movement fem/tm) was 2/5, 1/5, 1/10, 1/20, and 0. All these ratios were tested when the direction of the target movement was in the same (H+), opposite (H?), and at right angles to (V+, V?) the movement of the eyes. Eye movements, target movements, and reports of target movement were recorded. Results indicate that a discrepancy between eye and target movement greater than 20% predictably leads to perceived target movement, whereas a discrepancy of 5% or less rarely leads to perceived movement. The results are interpreted as support for the operation of a compensatory mechanism during eye movements.     

I know you are not looking at me: capuchin monkeys’ (<Emphasis Type="Italic">Cebus apella</Emphasis>) sensitivity to human attentional states

The present study asked whether capuchin monkeys recognize human attentional states. The monkeys requested food from the experimenter by extending an arm (pointing) toward the baited one of two transparent cups. On regular trials the experimenter gave the food immediately to the monkeys upon pointing but on randomly inserted test trials she ignored the pointing for 5 s during which she displayed different attentional states. The monkeys looked at the experimenter's face longer when she looked at the monkeys than when she looked at the ceiling in Experiment 1, and longer when she oriented her head midway between the two cups with eyes open than when she did so with eyes closed in Experiment 2. However, the monkeys showed no differential pointing in these conditions. These results suggest that capuchins are sensitive to eye direction but this sensitivity does not lead to differential pointing trained in laboratory experiments. Furthermore, to our knowledge, this is the first firm behavioral evidence that non-human primates attend to the subtle states of eyes in a food requesting task.     

Biases in eye movements to threatening facial expressions in generalized anxiety disorder and depressive disorder

The study investigated biases in selective attention to emotional face stimuli in generalized anxiety disorder (GAD) and depressive disorder, using a modified probe detection task. There were 4 face types: threatening, sad, happy, and neutral. Measures of attentional bias included (a) the direction and latency of the initial eye movement in response to the faces and (b) manual reaction time (RT) to probes replacing the face stimuli 1,000 ms after their onset. Results showed that individuals with GAD (without depressive disorder) were more likely to look first toward threat faces rather than neutral faces compared with normal controls and those with depressive disorder. They also shifted their gaze more quickly toward threat faces, rather than away from them, relative to the other two groups. There were no significant findings from the manual RT data. Implications of the results for recent theories of clinical anxiety and depression are discussed.