Some Aspects of the Motor Organization of the Oculomotor System

Oculomotor responses in looking to visual targets were examined for motor coordination parallel to that found in manual pointing. Ocular responses did not show any evidence of speed-accuracy tradeoff when the correlation of latency (reaction time) and accuracy was assessed. The oculomotor system did show a range effect, hidden under a general undershoot of target position and present only when no visual feedback about target position was available to allow correction during the movement. Cues to allow predictive timing of the response seemed very important for oculomotor reaction time. Processing of timing of movement may be handled differently for oculomotor than for manual orienting, and calculation of position in space may be processed in common, leading to both similarities and differences for the two motor systems.     

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 Orienting in Two-dimensional Space

This paper examines how the covert orienting of spatial attention affects motor responses to visual stimuli. Premotor theories, as well as hemi-field inhibition accounts of visual attention predict an increase in response times when a target stimulus appears in the opposite direction to a spatial cue. Some models also suggest that this meridional effect should be increased across oblique meridians. Two types of cue (central and peripheral) were used to orient attention towards locations prior to the onset of visual targets. Simple manual (press button) and saccadic responses were measured. No meridional effects were found with peripheral cues, whereas central cueing produced meridional effects across all meridians. Cueing effects did not vary significantly with two-dimensional axis for either manual or saccadic responses. Increases in response time with cue-target distance were found for both response and cue types. For saccades, distance gradients were shallower moving distally rather than proximally from the cued position. However, simple manual responses did not show this asymmetry. Orienting to central cues also modulated the amplitude of saccades. The results are consistent with an effect of attentional cues in oculomotor centres as well as the existence of actiondependent attentional representations. However, it is proposed that, rather than reflecting oculomotor programming, meridional effects arise from a directional organization within spatio-cognitive representations.     

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.     

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.     

Control of Rapid Arm Movements When Target Position Is Altered During Saccadic Suppression

This experiment examined whether rapid arm movements can be corrected in response to a change in target position that occurs just prior to movement onset, during saccadic suppression of displacement. Because the threshold of retinal input reaches its highest magnitude at that time, displacement of the visual target of a saccade is not perceived. Subjects (N = 6) were instructed to perform very rapid arm movements toward visual targets located 16, 20, and 24 degrees from midline (on average, movement time was 208 ms). On some trials the 20 degrees target was displaced 4 degrees either to the right or to the left during saccadic suppression. For double-step trials, arm movements did not deviate from their original trajectory. Movement endpoints and movement structure (i.e., velocity-and acceleration-time profiles) were similar whether or not target displacements occurred, showing the failure of proprioceptive signals or internal feedback loops to correct the arm trajectory. Following this movement, terminal spatially oriented movements corrected the direction of the initial movement (as compared with the single-step control trials) when the target eccentricity decreased by 4 degrees. Subjects were unaware of these spatial corrections. Therefore, spatial corrections of hand position were driven by the goal level of the task, which was updated by oculomotor corrective responses when a target shift occurred.     

Optimizing the Use of Vision in Manual Aiming: The Role of Practice

The purpose of this study was to determine how subjects learn to adjust the characteristics of their manual aiming movements in order to make optimal use of the visual information and reduce movement error. Subjects practised aiming (120 trials) with visual information available for either 400 msec or 600 msec. Following acquisition, they were transferred to conditions in which visual information was available for either more or less time. Over acquisition, subjects appeared to reduce target-aiming error by moving to the target area more quickly in order to make greater use of vision when in the vicinity of the target. With practice, there was also a reduction in the number of modifications in the movement. After transfer, both performance and kinematic data indicated that the time for which visual information was available was a more important predictor of aiming error than the similarity between training and transfer conditions. These findings are not consistent with a strong “specificity of learning” position. They also suggest that, if some sort of general representation or motor programme develops with practice, that representation includes rules or procedures for the utilization of visual feedback to allow for the on-line adjustment of the goal-directed movement.     

Changes in straight-ahead eye position during adaptation to wedge prisms

If S is instructed to look straight ahead before adapting to laterally displaced vision, he does so without noticeable error. After adapting, however, in response to the same instruction, he may rotate his eyes as much as 8° toward the the displaced visual target. This is the change in judgment of the direction of gaze which Helmholtz identified in 1867 as an important physiological mechanism in adaptation to prisms. It leads to more accurate reaching behavior by causing S to make a visual judgment that the target is closer to straight ahead than it was when he first looked through the prisms. This type of adaptive change (change in judgment of the direction of gaze, oculomotor change) can be measured either by manual judgments (difference between successive “straight ahead” and “visual target” judgments) or by changes in straight-ahead eye position. It may be described as a parametric adjustment in the oculomotor control system, and is closely analogous to the eye movement which subserves the recovery of binocular fusion in prism vergence.     

Effects of Visual Error Timing on Smooth Pursuit Gain Adaptation in Humans

Smooth pursuit (SP) is one of the precise oculomotor behaviors when tracking a moving object. Adaptation of SP is based on a visual-error driven motor learning process associated with predictable changes in the visual environment. Proper timing of a sensory signal is an important factor for adaptation of fine motor control. In this study, we investigated whether visual error timing affects SP gain adaptation. An adaptive change in SP gain is produced experimentally by repeated trials of a step-ramp tracking with 2 different velocities (double-velocity paradigm). The authors used the double-velocity paradigm where target speed changes 400 or 800 ms after the target onset. The results show that SP gain changed in a certain time window following adaptation. The authors suggest that SP adaptation shown in this study is associated with timing control mechanisms.     

Interference effects between saccadic and key-press reaction times of volleyball players and nonathletes

To investigate the interference effect in volleyball players and nonathletes (ns=10) when they executed both saccadic and key-press reaction time (RT) tasks concurrently, the two groups responded to the onset of peripheral visual stimuli as quickly as possible in single and dual conditions. In the single condition, subjects responded with either saccadic eye or key-press movement. In the dual condition, they responded concurrently with both saccadic eye and key-press movements. In both groups, the key-press RT was longer in the dual condition than in the single condition. However, the amount of key-press RT delay was remarkably smaller for the volleyball players than for nonathletes. This suggests the motor command to initiate manual movement of volleyball players might be less interfered with by a concurrent oculomotor command to initiate saccadic eye movement when compared to that of nonathletes.     

Target-size influences on reaction time with movement time controlled

The variable that affect motor programming time may be studied by changing the nature of the response and measuring the subsequent changes in reaction time (RT). One notion of motor programming suggests that aiming responses with reduced target size and/or increased target amplitude require more "complex" motor programs that require longer RTs. In a series of five experiments which movement time (MT) was experimentally varied target size neither influences RT when the movement amplitude was 2 or 30 cm nor when the target sizes differed by as much as a factor of 16:1. Increasing the movement amplitude from 15 to 30 cm also had no influence on RT. Movement time, however, did affect RT, with 200-msec movements having longer RTs than 120-msec movements. Target size and movement amplitude did not appear to be factors that influence programming time or program complexity.     

A visual representation and the control of manual aiming movements

Three experiments were conducted to determine if a representation of the movement environment is functional in the organization and control of limb movements, when direct visual contact with the environment is prevented. In Experiment 1, a visual rearrangement procedure was employed to show that a representation of the environment that provides inaccurate information about the spatial location of a target can disrupt manual target aiming. In Experiment 2, we demonstrated that spatial information about the position of a target can be destroyed by a visual pattern mask, supporting our claim that the representation is visual. A target-cuing procedure was used in Experiment 3 to show that representation of target position can be useful for premovement organization in a target-aiming task. Together our findings suggest that a short-lived visual representation of the movement environment may serve a useful role in the organization and control of limb movements.     

A Visual Representation and the Control of Manual Aiming Movements

Three experiments were conducted to determine if a representation of the movement environment is functional in the organization and control of limb movements, when direct visual contact with the environment is prevented. In Experiment 1, a visual rearrangement procedure was employed to show that a representation of the environment that provides inaccurate information about the spatial location of a target can disrupt manual target aiming. In Experiment 2, we demonstrated that spatial information about the position of a target can be destroyed by a visual pattern mask, supporting our claim that the representation is visual. A target-cuing procedure was used in Experiment 3 to show that representation of target position can be useful for premovement organization in a targetaiming task. Together our findings suggest that a short-lived visual representation of the movement environment may serve a useful role in the organization and control of limb movements.     

Against a role for attentional disengagement in the gap effect: A friendly amendment to Tam and Stelmach (1993)

Saccadic reaction time (RT) is reduced when the fixation point is removed shortly before target onset. Although Tam and Stelmach (1993) argued that thisgap effect could not be explained solely by the idea that fixation offset disengaged visual attention and preferred an explanation based on disengagement of the oculomotor system, they felt that they could not rule out a hybrid model in which both oculomotorand attentional disengagement contribute to the gap effect. Our analysis of the dual response experiment (Experiment 4), upon which this hybrid model was based, shows that manual and saccadic responses were likely compromised by a grouping or delay strategy and that subjects may not have been attending as instructed. On these grounds, we argue that Tam and Stelmach (1993), like Kingstone and Klein (1990; 1993a) provide no evidence that attentional disengagement contributes to the gap effect. An alternative proposal (Klein & Kingstone, 1993), that motor preparation and oculomotor disengagement combine additively to produce the gap effect, is consistent with the data from Tam and Stelmach’s Experiments 1–3, is similar to the explanation that they prefer, and has been strongly supported when directly tested (Kingstone, Klein, & Taylor, 1994).     

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.     

Timing and torque involvement in the organisation of a rapid forearm flexion

The study was designed to determine whether the magnitude of force and the timing of force are response parameters involved in the organisation of a rapid forearm flexion to a target. The magnitude of torque and the timing of torque were manipulated independently through manipulations of the total moment of inertia and movement time, and the effect of these manipulations on premotor and motor reaction times was observed. Planned comparison analyses revealed that premotor and motor reaction times increased when a movement, which required the same magnitude of torque as in a fast movement, was performed slower. However, premotor and motor reaction times were not affected when movements were performed at the same speed, but differed with respect to the magnitude of torque required. These results indicate that a different timing requirement in the forthcoming movement is associated with a corresponding change in the amount of central processing time required. Therefore, the timing of torque appears to be a parameter of the movement that is organised in advance of movement execution. However, a change in the specification for the magnitude of torque does not affect the amount of time needed to organise the movement.     

Lateralized effects of target location on reaction times when preparing for manual aiming at a visual target

To elucidate the temporal characteristics of information processing for motor action differing in complexity in relation to both perceptual and cognitive information processing, we investigated whether the reaction times (RTs) to a visual target would be affected by task complexity (finger lifting or manual aiming), pre-cueing (with a pre-cue or without a pre-cue), or target location (five horizontal positions). Using the right hand, seven right-handed subjects performed two tasks, finger lifting and manual aiming at a target, with or without a pre-cue. The pre-cue announced the location of the target to be presented. An ANOVA showed significant interactions between task and location and between pre-cue and location with no significant interaction between task and pre-cue, indicating that the task-location interaction does not depend on whether or not a pre-cue is given. The manual-aiming RTs were longer than the finger-lifting RTs, and the effects of the target location on the RTs differed for finger lifting and manual aiming. It can be assumed that the longer RTs of manual aiming reflect the time for information processing that is needed when preparing for the aiming action per se, which is an extra movement performed in addition to the simple initiation of finger lifting. Differential RTs (DRTs) calculated by subtracting the finger-lifting RTs from the aiming RTs were therefore examined. The DRTs significantly differed for target locations (i.e., a lateralized effect), with the DRTs for an ipsilateral target appearing to be significantly shorter than those for contralateral and central targets. The lateralized effect appearing on the DRTs may be mediated by the processing of visual-spatial information about visual targets as motor preparations are made for manual aiming.     

Saccade preparation inhibits reorienting to recently attended locations

We measured manual reaction time in normal human subjects to confirm that an eccentric visual signal has a biphasic effect on covert attention and eye movements. First, it summons attention and biases a saccade toward the signal; a subsequent inhibition of return then slows responses to signals at that location. A temporal hemifield dominance for inhibition of return was shown; this finding converges with observations in neurologic patients to suggest that it is mediated by midbrain pathways. Endogenous orienting of attention, from a central arrow cue, did not activate inhibition of return, whereas endogenous saccade preparation did so as effectively as an exogenous signal, even when no saccade was made. Inhibition of return is activated by midbrain oculomotor pathways and may function as a location "tagging" mechanism to optimize efficiency of visual search.     

Intermittency in human manual tracking tasks

We confirm Craik's (1947) observation that the human manual1y tracking a visual target behaves like an intermittent servo-control1er. Such tracking responses are indicative of "sampled" negative-feedback control but could be the result of other, continuous, mechanisms. Tracking performance therefore was recorded in a task in which visual feedback of the position of the hand-held joystick could be eliminated. Depriving the subjects of visual feedback led to smoother tracking and greatly reduced the signal power of their responses between 0.5-1.8 Hz. Their responses remained intermittent when they used feedback of their own position but not of the target to track a remembered (virtual) target. Hence, intermittency in tracking behavior is not exclusively a signature of visual feedback control but also may be a sign of feedback to memorized waveforms. Craik's (1947) suggestion that the intermittency is due to a refractory period following each movement was also tested. The errors measured at the start of each intermittent response, during tracking of slow waveforms, showed evidence of a small error deadzone (measuring 0.7 cm on the VDU screen or 0.80 degrees at the eye). At higher target speeds, however, the mean size of starting errors increased, and the upper boundary of the distribution of starting error was close to that expected of a refractory delay of approximately 170 ms between responses. We consider a model of the control system that can fit these results by incorporating an error deadzone within a feedback control loop. We therefore propose that the initiation of intermittent tracking responses may be limited by a positional error deadzone and that evidence for a refractory period between successive corrective movements can be satisfied without evoking an explicit timing or sampling mechanism.     

New insights on eye blindness and hand sight: Temporal constraints of visuo-motor networks

Pioneer experiments on saccadic suppression have shown that this effect is not followed by motor disorientation: Conscious perception of a target displacement can be dissociated from correct manual target reaching. It has subsequently been demonstrated that movement corrections with the same latency and spatial characteristics can be produced in conditions allowing perceptual awareness of perturbation of a target as in condition inducing saccadic suppression. In addition to the qualitative dissociation between motor performance and conscious awareness, quantitative temporal dissociations in action can be observed by manipulating different features of the visual target. When the target of an ongoing simple action is perturbed, a temporal advantage is found for responses to perturbations of location relative to colour and shape. Furthermore, there seems to be a temporal advantage for automatic motor corrections made in response to a target displacement as compared to other responses (other ongoing movement adjustments, movement interruption, conditional motor response such as pressing a key, verbal response, delayed matching-to-sample tasks). Thus, this paper reviews evidence for the fact that the temporal characteristics of any given response to a stimulus are dependent both on the sensory processes and on the type of response generated. Accordingly, identification responses (such as verbal report) to a visual stimulus are much slower than motor corrections of an ongoing movement in response to a target location change because of different processing times of the stimulus features (“What” compared to “Where”) and of the response itself (“What” compared to “How”). The existence of two continua (What/Where and What/How) is proposed between these two extreme stimulus- response combinations. This model may be a useful framework to better understand visuo-motor transformations and the network of connections between visual and motor areas.