Developmental changes in the control of saccadic eye movements in response to directional eye gaze and arrows

We investigated developmental differences in oculomotor control between 10-year-old children and adults using a central interference task. In this task, the colour of a fixation point instructed participants to saccade either to the left or to the right. These saccade directions were either congruent or incongruent with two types of distractor cue: either the direction of eye gaze of a centrally presented schematic face, or the direction of arrows. Children had greater difficulties inhibiting the distractor cues than did adults, which revealed itself in longer saccade latencies for saccades that were incongruent with the distractor cues as well as more errors on these incongruent trials than on congruent trials. Counter to our prediction, in terms of saccade latencies, both children and adults had greater difficulties inhibiting the arrow than the eye gaze distractors.     

Developmental changes in the control of saccadic eye movements in response to directional eye gaze and arrows

We investigated developmental differences in oculomotor control between 10-year-old children and adults using a central interference task. In this task, the colour of a fixation point instructed participants to saccade either to the left or to the right. These saccade directions were either congruent or incongruent with two types of distractor cue: either the direction of eye gaze of a centrally presented schematic face, or the direction of arrows. Children had greater difficulties inhibiting the distractor cues than did adults, which revealed itself in longer saccade latencies for saccades that were incongruent with the distractor cues as well as more errors on these incongruent trials than on congruent trials. Counter to our prediction, in terms of saccade latencies, both children and adults had greater difficulties inhibiting the arrow than the eye gaze distractors.     

The influence of eye-gaze and arrow pointing distractor cues on voluntary eye movements

We investigated Ricciardelli et al.'s (2002) claim, that the tendency for gaze direction to elicit automatic attentional following is unique to biologically significant information. Participants made voluntary saccades to targets on the left or the right of a display, which were either congruent or incongruent with a centrally presented distractor (eye-gaze or arrow). Contrary to Ricciardelli et al., for both distractor types, saccade latencies were slower, and participants made more directional errors, on incongruent than on congruent trials. Moreover, a cost-benefit analysis showed no difference between the two distractor types. However, latencies for erroneous saccades were faster than correctly directed saccades for the eye-gaze distractors, but not for the arrow distractors.     

Emotional attention: effects of emotion and gaze direction on overt orienting of visual attention

In the present study we considered the two factors that have been advocated for playing a role in emotional attention: perception of gaze direction and facial expression of emotions. Participants performed an oculomotor task in which they had to make a saccade towards one of the two lateral targets, depending on the colour of the fixation dot which appeared at the centre of the computer screen. At different time intervals (stimulus onset asynchronies, SOAs: 50,100,150 ms) following the onset of the dot, a picture of a human face (gazing either to the right or to the left) was presented at the centre of the screen. The gaze direction of the face could be congruent or incongruent with respect to the location of the target, and the expression could be neutral or angry. In Experiment 1 the facial expressions were presented randomly in a single block, whereas in Experiment 2 they were shown in separate blocks. Latencies for correct saccades and percentage of errors (saccade direction errors) were considered in the analyses. Results showed that incongruent trials determined a significantly higher percentage of saccade direction errors with respect to congruent trials, thus confirming that gaze direction, even when task-irrelevant, interferes with the accuracy of the observer’s oculomotor behaviour. The angry expression was found to hold attention for a longer time with respect to the neutral one, producing delayed saccade latencies. This was particularly evident at 100 ms SOA and for incongruent trials. Emotional faces may then exert a modulatory effect on overt attention mechanisms.     

Action effects in saccade control

According to the ideomotor principle, action preparation involves the activation of associations between actions and their effects. However, there is only sparse research on the role of action effects in saccade control. Here, participants responded to lateralized auditory stimuli with spatially compatible saccades toward peripheral targets (e.g., a rhombus in the left hemifield and a square in the right hemifield). Prior to the imperative auditory stimulus (e.g., a left tone), an irrelevant central visual stimulus was presented that was congruent (e.g., a rhombus), incongruent (e.g., a square), or unrelated (e.g., a circle) to the peripheral saccade target (i.e., the visual effect of the saccade). Saccade targets were present throughout a trial (Experiment 1) or appeared after saccade initiation (Experiment 2). Results showed shorter response times and fewer errors in congruent (vs. incongruent) conditions, suggesting that associations between oculomotor actions and their visual effects play an important role in saccade control.     

Gaze and arrow distractors influence saccade trajectories similarly

Perceiving someone's averted eye-gaze is thought to result in an automatic shift of attention and in the preparation of an oculomotor response in the direction of perceived gaze. Although gaze cues have been regarded as being special in this respect, recent studies have found evidence for automatic attention shifts with nonsocial stimuli, such as arrow cues. Here, we directly compared the effects of social and nonsocial cues on eye movement preparation by examining the modulation of saccade trajectories made in the presence of eye-gaze, arrows, or peripheral distractors. At a short stimulus onset asynchrony (SOA) between the distractor and the target, saccades deviated towards the direction of centrally presented arrow distractors, but away from the peripheral distractors. No significant trajectory deviations were found for gaze distractors. At the longer SOA, saccades deviated away from the direction of the distractor for all three distractor types, but deviations were smaller for the centrally presented gaze and arrow distractors. These effects were independent of whether line-drawings or photos of faces were used and could not be explained by differences in the spatial properties of the peripheral distractor. The results suggest that all three types of distractors (gaze, arrow, peripheral) can induce the automatic programming of an eye movement. Moreover, the findings suggest that gaze and arrow distractors affect oculomotor preparation similarly, whereas peripheral distractors, which are classically regarded as eliciting an automatic shift of attention and an oculomotor response, induce a stronger and faster acting influence on response preparation and the corresponding inhibition of that response.     

Asynchrony in discrete bimanual aiming: Evidence for visual strategies of coordination

The bimanual coupling literature supposes an inherent drive for synchrony between the upper limbs when making discrete bimanual movements. The level of synchrony is argued to be task dependent, reliant on the visual demands of the two targets, and the result of a complex pattern of hand and eye movements (Bingham, Hughes, & Mon-Williams, 2008 ; Riek, Tresilian, Mon-Williams, Coppard, & Carson, 2003 ). However, recent work by Bruyn and Mason ( 2009 ) suggests that temporal coordination is not solely influenced by visual saccades. In this experimental series, a total of 8 participants performed congruent movements to targets either near or far from the midline. Targets far from the midline, requiring a visual saccade, resulted in greater terminal asynchrony. Initial and terminal asynchrony were not consistent, but linked to the task demands at that stage of the movement. If the asynchrony evident at the end of a bimanual movement is due to a complex pattern of hand and eye movements then the removal of visual feedback should result in an increase in synchrony. Sixteen participants then completed congruent and incongruent bimanual aiming movements to near and/or far targets. Movements were made with or without visual feedback of hands and targets. Analyses revealed that movements made without visual feedback showed increased synchrony between the limbs, yet movements to incongruent targets still showed greater asynchrony. We suggest that visual constraints are not the sole cause of asynchrony in discrete bimanual movements.     

Anxiety and sensitivity to gaze direction in emotionally expressive faces

This study investigated the role of neutral, happy, fearful, and angry facial expressions in enhancing orienting to the direction of eye gaze. Photographs of faces with either direct or averted gaze were presented. A target letter (T or L) appeared unpredictably to the left or the right of the face, either 300 ms or 700 ms after gaze direction changed. Response times were faster in congruent conditions (i.e., when the eyes gazed toward the target) relative to incongruent conditions (when the eyes gazed away from the target letter). Facial expression did influence reaction times, but these effects were qualified by individual differences in self-reported anxiety. High trait-anxious participants showed an enhanced orienting to the eye gaze of faces with fearful expressions relative to all other expressions. In contrast, when the eyes stared straight ahead, trait anxiety was associated with slower responding when the facial expressions depicted anger. Thus, in anxiety-prone people attention is more likely to be held by an expression of anger, whereas attention is guided more potently by fearful facial expressions.     

Affective priming of emotional pictures in parafoveal vision: Left visual field advantage

This study investigated whether stimulus affective content can be extracted from visual scenes when these appear in parafoveal locations of the visual field and are foveally masked, and whether there is lateralization involved. Parafoveal prime pleasant or unpleasant scenes were presented for 150 msec 2.5° away from fixation and were followed by a foveal probe scene that was either congruent or incongruent in emotional valence with the prime. Participants responded whether the probe was emotionally positive or negative. Affective priming was demonstrated by shorter response latencies for congruent than for incongruent prime-probe pairs. This effect occurred when the prime was presented in the left visual field at a 300-msec prime-probe stimulus onset asynchrony, even when the prime and the probe were different in physical appearance and semantic category. This result reveals that the affective significance of emotional stimuli can be assessed early through covert attention mechanisms, in the absence of overt eye fixations on the stimuli, and suggests that right-hemisphere dominance is involved.     

Arrows don’t look at you: Qualitatively different attentional mechanisms triggered by gaze and arrows

Eye gaze conveys rich information concerning the states of mind of others, playing a critical role in social interactions, signaling internal states, and guiding others’ attention. On the basis of its social significance, some researchers have proposed that eye gaze may represent a unique attentional stimulus. However, contrary to this notion, the majority of the literature has shown indistinguishable attentional effects when eye gaze and arrows have been used as cues. Taking a different approach, in this study we aimed at finding qualitative attentional differences between gazes and arrows when they were used as targets instead of as cues. We used a spatial Stroop task, in which participants were required to identify the direction of eyes or arrows presented to the left or the right of a fixation point. The results showed that the two types of stimuli led to opposite spatial interference effects, with arrows producing faster reaction times when the stimulus direction was congruent with the stimulus position (a typical spatial Stroop effect), and eye gaze producing faster reaction times when it was incongruent (a “reversed” spatial Stroop effect). This reversed Stroop is interpreted as an eye-contact effect, therefore revealing the unique nature of eyes as special social-attention stimuli.     

Effects of saccades and response type on the Simon effect: If you look at the stimulus,the Simon effect may be gone

The Simon effect has most often been investigated with key-press responses and eye fixation. In the present study, we asked how the type of eye movement and the type of manual response affect response selection in a Simon task. We investigated three eye movement instructions (spontaneous, saccade, and fixation) while participants performed goal-directed (i.e., reaching) or symbolic (i.e., finger-lift) responses. Initially, no oculomotor constraints were imposed, and a Simon effect was present for both response types. Next, eye movements were constrained. Participants had to either make a saccade toward the stimulus or maintain gaze fixed in the screen centre. While a congruency effect was always observed in reaching responses, it disappeared in finger-lift responses. We suggest that the redirection of saccades from the stimulus to the correct response location in noncorresponding trials contributes to the Simon effect. Because of eye–hand coupling, this occurred in a mandatory manner with reaching responses but not with finger-lift responses. Thus, the Simon effect with key-presses disappears when participants do what they typically do—look at the stimulus.     

Endogenous saccades are preceded by shifts of visual attention: evidence from cross-saccadic priming effects

The present study examines whether endogenous saccades are preceded by shifts of attention. Three experiments are reported in which participants were required to execute a saccadic eye movement to a certain location and to subsequently identify the orientation of a target triangle. Prior to the execution of the saccade a prime was presented, which was compatible or incompatible with the target. A priming effect (faster responses in the compatible condition than in the incompatible condition) occurred only when the prime was presented at the saccade destination, and this effect was larger when the prime was presented during oculomotor programming than when it was presented prior to oculomotor programming. The results indicate that an endogenous shift of attention precedes endogenous saccades, providing further support for theories of visual selection that assume a tight coupling between attention and saccades.     

Age-related differences in automatic stimulus-response associations: Insights from young and older adults’ parity judgments

Young and older adults completed a parity judgment task (i.e., judging whether a target digit was odd or even) in which target numbers were preceded by masked prime numbers presented for 43 msec. Targets were either congruent (i.e., they had the same parity status as their primes) or incongruent (i.e., odd primes were paired with even targets, and even primes were paired with odd targets). Response times, percent errors, and event-related potentials (ERPs) were recorded for all items to compare automatic stimulus—response association (ASRA) and congruence effects (i.e., better performance on congruent than on incongruent trials) across age groups. Two important original sets of findings were obtained in this sample of participants. First, both age groups showed ASRA effects in behavioral measures. Second, age-related differences were observed in amplitude, timing, and scalp distributions for each congruent and incongruent ERP. These findings have implications for furthering the understanding of ASRA effects and of general characteristics of cognitive processes affected (or not affected) by aging.     

Response inhibition in motor and oculomotor conflict tasks: Different mechanisms,different dynamics?

Previous research has shown that the appearance of task-irrelevant abrupt onsets influences saccadic eye movements during visual search and may slow down manual reactions to target stimuli. Analysis of reaction time distributions in the present study offers evidence suggesting that top-down inhibition processes actively suppress oculomotor or motor responses elicited by a salient distractor, in order to resolve the conflict that arises when reflex-like and deliberate responses are in opposition. Twenty-six participants carried out a variation of the oculomotor capture task. They were instructed to respond with either a saccade toward or with a button press at the side of the hemifield in which a target color singleton appeared. A distractor stimulus could appear either in the same or in the opposite hemifield. Delta plots revealed competition between reflex-like and deliberate response activation, and highlighted selective inhibition of automatic responses: While participants generally responded more slowly in incongruent compared to congruent situations, this effect diminished and even reversed in the slowest speed quantiles. These effects were present in both the oculomotor and motor response-mode conditions.     

The effect of blinks and saccadic eye movements on visual reaction times

Vision is suppressed during blinks and saccadic eye movements. We hypothesized that visual reaction times (RTs) in a vigilance test would be significantly increased when a blink or a saccade happened to coincide with the stimulus onset. Thirty healthy volunteers each performed a visual RT test for 15 min while their eye and eyelid movements were monitored by a system of infrared reflectance oculography. RTs increased significantly, many by more than 200 msec, when a blink occurred between 75 msec before and up to 150 msec after the stimulus onset. A similar result was observed with saccades that started 75 to 150 msec after the stimulus. Vision or attention was evidently inhibited before each blink and for longer than the saccades lasted. We suggest that visual suppression is involved in this process, which could explain some of the normal variability in RTs over periods of seconds that has not been adequately explained before.     

Does gaze perception facilitate overt orienting?

Recent studies have demonstrated the facilitation of responses to peripheral targets cued by the direction of the gaze of a face. However, in the absence of data from an eye tracker, it has been unclear to what extent these effects are due to the participants making small saccades in response to the cue that bring them closer to the congruent location of the cued targets. We used an eye tracker to show that while such cue-driven saccades occur, they do not account for the main cueing effects observed. Additionally, by using the same general paradigm as has previously been used with infants, we suggest that different mechanisms underlie eye gaze cueing effects in infants and adults.     

Gaze cueing as a function of perceived gaze direction

Previous studies have found that attention is automatically oriented in the direction of other people's gaze. This study directly investigated whether the perceiving gaze direction modulates the orienting of observers' attention. Gaze perception was manipulated by changing the face context (head orientation) of the gaze cue: the perceived gaze angle was increased (or decreased) when the head and gaze are congruent (or incongruent), while the local‐feature information of the eye region was preserved for all stimuli. The results showed that gaze‐cueing effects were enhanced when the perceived gaze direction was averted more toward left or right, and reduced when the perceived gaze direction was closer to direct gaze. The results suggest that gaze‐cueing effects are based on mechanisms specialized for gaze perception, and the magnitude of gaze‐cueing effects was probably a function of the perceived gaze direction.     

The influence of head contour and nose angle on the perception of eye-gaze direction

We report seven experiments that investigate the influence that head orientation exerts on the perception of eye-gaze direction. In each of these experiments, participants were asked to decide whether the eyes in a brief and masked presentation were looking directly at them or were averted. In each case, the eyes could be presented alone, or in the context of congruent or incongruent stimuli In Experiment 1A, the congruent and incongruent stimuli were provided by the orientation of face features and head outline. Discrimination of gaze direction was found to be better when face and gaze were congruent than in both of the other conditions, an effect that was not eliminated by inversion of the stimuli (Experiment 1B). In Experiment 2A, the internal face features were removed, but the outline of the head profile was found to produce an identical pattern of effects on gaze discrimination, effects that were again insensitive to inversion (Experiment 2B) and which persisted when lateral displacement of the eyes was controlled (Experiment 2C). Finally, in Experiment 3A, nose angle was also found to influence participants' ability to discriminate direct gaze from averted gaze, but here the effect was eliminated by inversion of the stimuli (Experiment 3B). We concluded that an image-based mechanism is responsible for the influence of head profile on gaze perception, whereas the analysis of nose angle involves the configural processing of face features.     

Covert orienting of attention in 3-month-old infants: The case of biological motion

Humans attend to different positions in the space either by moving their eyes or by moving covertly their attention. The development of covert attention occurs during the first year of life. According to Colombo’s model of attention (2001), within the first years there is a significant change in infants’ visuo-spatial orienting mechanisms, from a predominantly overt form to a covert orienting starting from 4 to 5 months of life. The use of non-invasive brain imaging techniques can shed light on the origin of such mechanisms. In particular, EEG and ERP studies can directly investigate the neural correlates of covert attention in young infants. The present study investigated the neural correlates of covert attention employing a visuo-spatial cueing paradigm in 3-month-old infants. Infants were presented with a central point-light walker (PLD) followed by a single peripheral target. The target appeared randomly at a position either congruent or incongruent with the walking direction of the cue. We examined infants’ target-locked P1 component and the saccade latencies toward the peripheral target. Results showed that the P1 component was larger in response to congruent than to incongruent targets and saccade latencies were faster for congruent rather than incongruent trials. Moreover, the facilitation in processing sensory information (priming effects) presented at the cued spatial location occurs even before the onset of the oculomotor response, suggesting that covert attention is present before 4 months of age. Overall, this study highlights how ERPs method could help researchers at investigating the neural basis of attentional mechanisms in infants.     

Saccadic Eye Movements in Parkinson's Disease: I. Delayed Saccades

The saccadic eye movements of nine patients with Parkinson's disease were compared to those of nine age-matched controls in two paradigms generating volitional saccades. In both paradigms, subjects had to make delayed saccades to peripheral LED targets: a peripheral target appeared 700 msec before a buzzer sounded, the buzzer being the signal to make a saccade to the target. In the first paradigm (“centre-off”), the fixation target was extinguished simultaneously with buzzer onset. In the second (“centre-remain”) it was not extinguished until 1000 msec later. The results showed that for outward saccades in both paradigms, there was no difference between Parkinsonian patients and controls, but saccadic latencies were significantly shorter in the “centre-remain” paradigm. The initial outward saccades were indistinguishable from the normal, reflex saccades of the same subjects. However, saccades returning to the centre (a type of remembered target saccade) were hypometric and showed multistepping. Both effects were more pronounced in patients with Parkinson's disease. The significance of these findings in terms of current hypotheses about the nature of the Parkinsonian saccadic deficit is discussed.