Impaired initiation but not execution of contralesional saccades in hemispatial neglect

Patients with unilateral neglect are impaired at making saccades to contralesional targets. Whether this problem arises from a deficit in perception, in planning the saccade or in executing the eye movement or some combination thereof remains unclear. We measured several variables related to the initiation and execution of saccades in an experiment which crossed two factors: target side (left, right) and direction of saccade (leftwards, rightwards). Relative to control subjects, patients with left-sided neglect were impaired in planning but not executing the contralesional saccade; while the latency to move their eyes following the onset of the target was increased, the duration and velocity to reach the target were normal. In addition, there were also no directional differences for saccades that were hypometric or inaccurate in the patients, further ruling out an execution impairment. Interestingly, this directional initiation deficit was exaggerated for leftward saccades to left targets, compared with all other conditions. We suggest that the disadvantage for contralesional saccades in neglect patients is attributable to a deficit not only in perceiving contralateral targets but also in planning leftward saccades. Once the saccade is initiated, however, execution apparently proceeds unimpaired.     

Lexical processing during saccades in text comprehension

We asked whether people process words during saccades when reading sentences. Irwin (1998) demonstrated that such processing occurs when words are presented in isolation. In our experiment, participants read part of a sentence ending in a high- or low-frequency target word and then made a long (40°) or short (10°) saccade to the rest of the sentence. We found a frequency effect on the target word and the first word after the saccade, but the effect was greater for short than for long saccades. Readers therefore performed more lexical processing during long saccades than during short ones. Hence, lexical processing takes place during saccades in text comprehension.     

Distractors that signal reward attract the eyes

Salient stimuli and stimuli associated with reward have the ability to attract both attention and the eyes. The current study exploited the effects of reward on the well-known global effect in which two objects appear simultaneously in close spatial proximity. Participants always made saccades to a predefined target, while the colour of a nearby distractor signalled the reward available (high/low) for that trial. Unlike previous reward studies, in the current study these distractors never served as targets. We show that participants made fast saccades towards the target. However, saccades landed significantly closer to the high compared to the low reward signalling distractor. This reward effect was already present in the first block and remained stable throughout the experiment. Instead of landing exactly in between the two stimuli (i.e., the classic global effect), the fastest eye movements landed closer towards the reward signalling distractor. Results of a control experiment, in which no distractor-reward contingencies were present, confirmed that the observed effects were driven by reward and not by physical salience. Furthermore, there were trial-by-trial reward priming effects in which saccades landed significantly closer to the high instead of the low reward signalling distractor when the same distractor was presented on two consecutive trials. Together the results imply that a reward signalling stimulus that was never part of the task set has an automatic effect on the oculomotor system.     

Saccades to targets in three-dimensional space: dependence of saccadic latency on target location.

The latency of saccadic movements to targets appearing at various positions in three-dimensional visual space was measured in four experiments. The first experiment confirmed that latencies of saccades to visual targets are greater in the lower visual field and showed that the increase is not influenced by the vertical starting position of the eye in the orbit, nor by a time gap between the fixation offset and the target onset. A hypothesis that this visual field difference was caused by a link between downward saccades and convergence movements was tested by recording saccade latencies when the targets were in a different depth plane from that of the original fixation. We did not find any direct support for the vergence involvement hypothesis, although the lower/upper visual field effect was shown to decrease consistently in monocular viewing. It was also shown that saccades to targets positioned in a different depth plane have longer latencies. In a final experiment, the visual field effect was shown to depend on the egocentric rather than the gravitational vertical.     

Saccades to targets in three-dimensional space: Dependence of saccadic latency on target location

The latency of saccadic movements to targets appearing at various positions in three-dimensional visual space was measured in four experiments. The first experiment confirmed that latencies of saccades to visual targets are greater in the lower visual field and showed that the increase is not influenced by the vertical starting position of the eye in the orbit, nor by a time gap between the fixation offset and the target onset. A hypothesis that this visual field difference was caused by a link between downward saccades and convergence movements was tested by recording saccade latencies when the targets were in a different depth plane from that of the original fixation. We did not find any direct support for the vergence involvement hypothesis, although the lower/upper visual field effect was shown to decrease consistently in monocular viewing. It was also shown that saccades to targets positioned in a different depth plane have longer latencies. In a final experiment, the visual field effect was shown to depend on the egocentric rather than the gravitational vertical.     

The Role of Emotional Content and Perceptual Saliency During the Programming of Saccades Toward Faces

Previous studies have shown that the human visual system can detect a face and elicit a saccadic eye movement toward it very efficiently compared to other categories of visual stimuli. In the first experiment, we tested the influence of facial expressions on fast face detection using a saccadic choice task. Face-vehicle pairs were simultaneously presented and participants were asked to saccade toward the target (the face or the vehicle). We observed that saccades toward faces were initiated faster, and more often in the correct direction, than saccades toward vehicles, regardless of the facial expressions (happy, fearful, or neutral). We also observed that saccade endpoints on face images were lower when the face was happy and higher when it was neutral. In the second experiment, we explicitly tested the detection of facial expressions. We used a saccadic choice task with emotional-neutral pairs of faces and participants were asked to saccade toward the emotional (happy or fearful) or the neutral face. Participants were faster when they were asked to saccade toward the emotional face. They also made fewer errors, especially when the emotional face was happy. Using computational modeling, we showed that this happy face advantage can, at least partly, be explained by perceptual factors. Also, saccade endpoints were lower when the target was happy than when it was fearful. Overall, we suggest that there is no automatic prioritization of emotional faces, at least for saccades with short latencies, but that salient local face features can automatically attract attention.     

Presaccadic attention allocation and express saccades

Express saccades are visually-guided saccades that are characterized by an extremely short latency of about 100 ms. The present experiments tested the hypothesis that a disengagement of visual attention is necessary for the generation of express saccades. All subjects produced large numbers of express saccades in the gap paradigm, in which the fixation stimulus is removed 200 ms before target onset (Exp. 1), but not in the overlap paradigm, in which the fixation stimulus remains on during the entire trial (Exp. 2). By means of peripheral cues (Exps. 3–5) and central cues (Exps. 6–7), visual attention was directed at the target location for the saccade before the actual appearance of the saccade target. In all experiments, the location cues facilitated rather than abolished express saccades. The generation of express saccades was facilitated even when the currently fixated visual stimulus was not removed before target onset (fixation-overlap; Exps. 5–7). The results are explained by the hypothesis that a disengagement of a separate fixation system is necessary for the generation of express saccades, a hypothesis that is in line with current neurobiological findings.     

Saccadic suppression of displacement: effects of illumination and background manipulation

In contrast to other functions which are suppressed during saccades, saccadic suppression of displacement (SSD--a decrease in sensitivity to visual displacements during saccades) has often been considered to be due to efferent processes rather than to visual masking. The aim of this study was to explicitly assess the importance of visual conditions in SSD. In two experiments, a small computer-generated target made random horizontal jumps. An infrared eye tracker was used to detect the saccade toward the new position, triggering a smaller centripetal displacement of the target. Subjects reported awareness of these intrasaccadic displacements by pressing a key. In the first experiment, the task was performed in both a well-lit environment and in darkness. In the second experiment these conditions were replicated and additional factors such as the contrast of the background and the effect of moving the target spot alone or the target plus the entire background were investigated. Unlike other forms of saccadic suppression, SSD was stronger in the dark, although subjects also had a greater bias to report detections in that condition. Other background manipulations had no effect. The effect of ambient lighting on SSD is small and subtle. Effects of other background manipulations may be overridden by the focusing of attention on a small moving target.     

Visual search is modulated by action intentions

The influence of action intentions on visual selection processes was investigated in a visual search paradigm. A predefined target object with a certain orientation and color was presented among distractors, and subjects had to either look and point at the target or look at and grasp the target. Target selection processes prior to the first saccadic eye movement were modulated by the different action intentions. Specifically, fewer saccades to objects with the wrong orientation were made in the grasping condition than in the pointing condition, whereas the number of saccades to an object with the wrong color was the same in the two conditions. Saccadic latencies were similar under the different task conditions, so the results cannot be explained by a speed-accuracy trade-off. The results suggest that a specific action intention, such as grasping, can enhance visual processing of action-relevant features, such as orientation. Together, the findings support the view that visual attention can be best understood as a selection-for-action mechanism.     

The planning and execution of sequential eye movements: saccades do not show the one target advantage

The present experiment examined the one-target advantage (OTA) with regard to saccadic eye movements. The OTA, previously found with manual pointing responses, refers to the finding that movements are executed faster when the limb is allowed to stop on the target compared to the situation where it has to proceed and hit a second target. Using an adapted limb movement OTA task, saccades of 5 degrees and 15 degrees were made to (a) a single target (one-target), (b) one target and immediately to another target without a change in direction (two-target-extension), and (c) one target and immediately back to the start location (two-target-reversal). Unlike manual movements, the movement times for the initial saccade in the two-target-extension condition were not prolonged compared to either of the other two conditions. Moreover, this pattern of results was found for both the shorter and longer amplitude saccades. The results indicate that the OTA does not occur in the oculomotor system and therefore is not a general motor control phenomenon.     

EGOCENTRIC ACTION IN EARLY INFANCY:

Abstract— The extent to which infants combine visual (i e, retinal position) and nonvisual (eye or head position) spatial information in planning saccades relates to the issue of what spatial frame or frames of reference influence early visually guided action. We explored this question by testing infants from 4 to 6 months of age on the double-step saccade paradigm, which has shown that adults combine visual and eye position information into an egocentric (head- or trunk-centered) representation of saccade target locations. In contrast, our results imply that infants depend on a simple retinocentric representation at age 4 months, but by 6 months use egocentric representations more often to control saccade planning. Shifts in the representation of visual space for this simple sensorimotor behavior may index maturation in cortical circuitry devoted to visual spatial processing in general.     

A new look at the attribution of moral responsibility: The underestimated relevance of social roles

What are the main features that influence our attribution of moral responsibility? It is widely accepted that there are various factors which strongly influence our moral judgments, such as the agent’s intentions, the consequences of the action, the causal involvement of the agent, and the agent’s freedom and ability to do otherwise. In this paper, we argue that this picture is incomplete: We argue that social roles are an additional key factor that is radically underestimated in the extant literature. We will present an experiment to support this claim.     

Suppressing where but not what: the effect of saccades on dorsal- and ventral-stream visual processing

Some cognitive processes are suppressed during saccadic eye movements, whereas others are not. In two experiments, we investigated the locus of this interference effect. In one experiment, subjects decided whether pictured items were objects or nonobjects while making saccades of different lengths. Saccade distance had no effect on response time, indicating that saccades do not interfere with object recognition. However, in a second experiment, in which subjects decided whether pictured items faced to the left or to the right, response time increased with saccade distance, indicating that processing was suppressed during the saccade. These results (along with others) suggest that dorsal-stream (where) processes are suppressed during saccades, whereas ventral-stream (what) processes are not. Because the dorsal stream is instrumental in generating saccades, we propose that cognitive saccadic suppression results from dual-task interference within this visual subsystem.     

Visual perception of direction when voluntary saccades occur. I. Relation of visual direction of a fixation target extinguished before a saccade to a flash presented during the saccade

In experiments designed to clarify the mechanisms underlying the normal stability of visual direction for stationary objects when voluntary saccades occur, Ss reported on the horizontal visual direction of a brief test [lash presented when the eye was at a specific point in the saccade (the trigger point) relative to a fixation target viewed and extinguished prior to the saccade. From these reports, PSEs (points of subjective equality) were calculated for the fixation target as measured by the test [lashes. The distance of the trigger point from the previous fixation position was systematically varied in each experiment. Different experiments required saccades of different lengths and directions. With the exception of the presentation of the test [lash the saccades were carried out in complete darkness so that the possible utilization of an extraretinal signal regarding the eye movement (change in eye position, the intention to turn the eye, or a change of attention related to the eye movement) in the determination of visual direction could be observed uncomplicated by a continuing visual context. According to classical theories, an extraretinal signal proportional to the change in eye position acts to maintain direction constancy by compensating for the Shift of the retinal image resulting from the movement of the eye. In general, direction constancy was not preserved in the present experiments, and thus the data would not be predicted by classical theories. However, the PSE varied with distance of the trigger point from the fixation target. Since this displacement of PSE from the trigger point was in the correct direction for compensation, the presence of an extraretinal signal was confirmed. However, the growth of this signal appears to be time-locked to the saccade rather than locked to eye position; it is suggested that this growth takes place over a time period which is longer than the duration of the saccade itself.     

Programming saccadic eye movements

This article addresses questions about the preparatory processes that immediately precede saccadic eye movements. Saccade latencies were measured in a task in which subjects were provided partial advance information about the spatial location of a target fixation. In one experiment, subjects were faster in initiating saccades when they knew either the direction or amplitude of the required movement in advance compared to a condition with equal uncertainty about the number of potential saccade targets but without knowledge of the parameters required to execute the movement. These results suggest that the direction and amplitude for an upcoming saccade were calculated separately, and not in a fixed serial order. In another experiment, subjects appear to have programmed the saccades more holistically--with computations of direction and amplitude parameters occurring simultaneously. The implications of these results for models of eye movement preparation are discussed.     

Growing older does not always mean moving slower: examining aging and the saccadic motor system

Although humans typically move more slowly as they age, one exception may be the saccadic motor system. To fully determine whether the execution of saccades is affected by age, the authors examined detailed kinematics of vertical and horizontal saccades across a range of saccadic amplitudes (4 degrees, 8 degrees, and 12 degrees). Ten younger and 20 older adults participated in each experiment. Whereas in the 1st experiment, the authors assessed volitionally generated saccades, in the 2nd experiment, they evaluated reflexively generated saccades. The results of those experiments showed that the saccadic motor system is relatively impervious to the effects of aging; in fact, the differences between vertical and horizontal saccades were more evident than were differences between saccades produced by younger and older adults. The authors discuss possible reasons for that relative resistance to aging.     

Object files across eye movements: Previous fixations affect the latencies of corrective saccades

One of the factors contributing to a seamless visual experience is object correspondence—that is, the integration of pre- and postsaccadic visual object information into one representation. Previous research had suggested that before the execution of a saccade, a target object is loaded into visual working memory and subsequently is used to locate the target object after the saccade. Until now, studies on object correspondence have not taken previous fixations into account. In the present study, we investigated the influence of previously fixated information on object correspondence. To this end, we adapted a gaze correction paradigm in which a saccade was executed toward either a previously fixated or a novel target. During the saccade, the stimuli were displaced such that the participant’s gaze landed between the target stimulus and a distractor. Participants then executed a corrective saccade to the target. The results indicated that these corrective saccades had lower latencies toward previously fixated than toward nonfixated targets, indicating object-specific facilitation. In two follow-up experiments, we showed that presaccadic spatial and object (surface feature) information can contribute separately to the execution of a corrective saccade, as well as in conjunction. Whereas the execution of a corrective saccade to a previously fixated target object at a previously fixated location is slowed down (i.e., inhibition of return), corrective saccades toward either a previously fixated target object or a previously fixated location are facilitated. We concluded that corrective saccades are executed on the basis of object files rather than of unintegrated feature information.     

Saccadic suppression relies on luminance information

To determine whether saccadic suppression of image displacement uses information from luminance channels, we measured spatial displacement detection thresholds with equiluminant and nonequiluminant targets during saccades. We compared these saccadic thresholds with displacement thresholds measured during fixation by making ratios of saccadic thresholds to fixation thresholds. Ratios were lower in the equiluminant condition than in the nonequiluminant. This surprising result indicates that detection of equiluminant target displacements during saccades was better than detection of nonequiluminant targets, compared with the detection abilities during fixation. Thus, saccadic suppression of image displacement, which should increase displacement thresholds during saccades over fixation thresholds, was more effective with nonequiluminant targets. Because of target flicker, displacement thresholds were anisotropic in the nonequiluminant condition; thresholds were greater when target and eye moved in the same direction than when they moved in opposite directions, consistent with earlier results. These two effects (flicker-induced anisotropy and greater suppression in nonequiluminance) canceled when the eye moved opposite the displacement, yielding equal thresholds, and summed when eye and target moved in the same direction, yielding large threshold differences. We conclude that saccadic suppression of image displacement uses mechanisms sensitive to luminance contrast.     

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.     

Target localization after saccades and at fixation: Nontargets both facilitate and bias responses

ABSTRACT

The image on our retina changes every time we make an eye movement. To maintain visual stability after saccades, specifically to locate visual targets, we may use nontarget objects as “landmarks”. In the current study, we compared how the presence of nontargets affects target localization after saccades and during sustained fixation. Participants fixated a target object, which either maintained its location on the screen (sustained-fixation trials), or displaced to trigger a saccade (saccade trials). After the target disappeared, participants reported the most recent target location with a mouse click. We found that the presence of nontargets decreased response error magnitude and variability. However, this nontarget facilitation effect was not larger for saccade trials than sustained-fixation trials, indicating that nontarget facilitation might be a general effect for target localization, rather than of particular importance to post-saccadic stability. Additionally, participants’ responses were biased towards the nontarget locations, particularly when the nontarget-target relationships were preserved in relative coordinates across the saccade. This nontarget bias interacted with biases from other spatial references, e.g., eye movement paths, possibly in a way that emphasized non-redundant information. In summary, the presence of nontargets is one of several sources of reference that combine to influence (both facilitate and bias) target localization.