A simple rule of thumb for elegant prehension.

Reaching out to grasp an object (prehension) is a deceptively elegant and skilled behavior. The movement prior to object contact can be described as having two components, the movement of the hand to an appropriate location for gripping the object, the "transport" component, and the opening and closing of the aperture between the fingers as they prepare to grip the target, the "grasp" component. The grasp component is sensitive to the size of the object, so that a larger grasp aperture is formed for wider objects; the maximum grasp aperture (MGA) is a little wider than the width of the target object and occurs later in the movement for larger objects. We present a simple model that can account for the temporal relationship between the transport and grasp components. We report the results of an experiment providing empirical support for our "rule of thumb." The model provides a simple, but plausible, account of a neural control strategy that has been the center of debate over the last two decades.

The role of the primary motor cortex during skill acquisition on a two-degrees-of-freedom movement task.

One can partially eliminate motor skills acquired through practice in the hours immediately following practice by applying repetitive transcranial stimulation (rTMS) over the primary motor cortex. The disruption of acquired levels of performance has been demonstrated on tasks that are ballistic in nature. The authors investigated whether motor recall on a discrete aiming task is degraded following a disruption of the primary motor cortex induced via rTMS. Participants (N = 16) maintained acquired performance levels and patterns of muscle activity following the application of rTMS, despite a reduction in corticospinal excitability. Disruption of the primary motor cortex during a consolidation period did not influence the retention of acquired skill in this type of discrete visuomotor task.

Effects of acoustic startle stimuli on interceptive action.

In reaction time (RT) tasks, presentation of a startling acoustic stimulus (SAS) together with a visual imperative stimulus can dramatically reduce RT while leaving response execution unchanged. It has been suggested that a prepared motor response program is triggered early by the SAS but is not otherwise affected. Movements aimed at intercepting moving targets are usually considered to be similarly governed by a prepared program. This program is triggered when visual stimulus information about the time to arrival of the moving target reaches a specific criterion. We investigated whether a SAS could also trigger such a movement. Human experimental participants were trained to hit moving targets with movements of a specific duration. This permitted an estimate of when movement would begin (expected onset time). Startling and sub-startle threshold acoustic probe stimuli were delivered unexpectedly among control trials: 65, 85, 115 and 135 ms prior to expected onset (10:1 ratio of control to probe trials). Results showed that startling probe stimuli at 85 and 115 ms produced early response onsets but not those at 65 or 135 ms. Sub-threshold stimuli at 115 and 135 ms also produced early onsets. Startle probes led to an increased vigor in the response, but sub-threshold probes had no detectable effects. These data can be explained by a simple model in which preparatory, response-related activation builds up in the circuits responsible for generating motor commands in anticipation of the GO command. If early triggering by the acoustic probes is the mechanism underlying the findings, then the data support the hypothesis that rapid interceptions are governed by a motor program.

Triggering prepared actions by sudden sounds: reassessing the evidence for a single mechanism.

Loud acoustic stimuli can unintentionally elicit volitional acts when a person is in a state of readiness to execute them (the StartReact effect). It has been assumed that the same subcortical pathways and brain regions underlie all instances of the StartReact effect. They are proposed to involve the startle reflex pathways, and the eliciting mechanism is distinct from other ways in which sound can affect the motor system. We present an integrative review which shows that there is no evidence to support these assumptions. We argue that motor command generation for learned, volitional orofacial, laryngeal and distal limb movements is cortical and the StartReact effect for such movements involves transcortical pathways. In contrast, command generation for saccades, locomotor corrections and postural adjustments is subcortical and subcortical pathways are implicated in the StartReact effect for these cases. We conclude that the StartReact effect is not a special phenomenon mediated by startle reflex pathways, but rather is a particular manifestation of the excitatory effects of intense stimulation on the central nervous system.

Developmental changes in the response to obstacles during prehension.

Adults are proficient at reaching to grasp objects of interest in a cluttered workspace. The issue of concern, obstacle avoidance, was studied in 3 groups of young children aged 11-12, 9-10, and 7-8 years (n=6 in each) and in 6 adults aged 18-24 years. Adults slowed their movements and decreased their maximum grip aperture when an obstacle was positioned close to a target object (the effect declined as the distance between target and obstacle increased). The children showed the same pattern, but the magnitude of the effect was quite different. In contrast to the adults, the obstacle continued to have a large effect when it was some distance from the target (and provided no physical obstruction to movement).

Increasing confidence in vergence as a cue to distance.

Multiple cues contribute to the visual perception of an object's distance from the observer. The manner in which the nervous system combines these various cues is of considerable interest. Although it is accepted that image cues play a significant role in distance perception, controversy exists regarding the use of kinaesthetic information about the eyes' state of convergence. We used a perturbation technique to explore the contribution of vergence to visually based distance estimates as a function of both fixation distance and the availability of retinal information. Our results show that the nervous system increases the weighting given to vergence as (i) fixation distance becomes closer; and (ii) the available retinal image cues decrease. We also identified the presence of a strong contraction bias when distance cues were studied in isolation, but we argue that such biases do not suggest that vergence provides an ineffectual signal for near-space perception.

Improving vision: neural compensation for optical defocus.

Anecdotal reports abound of vision improving in myopia after a period of time without refractive correction. We explored whether this effect is due to an increased tolerance of blur or whether it reflects a genuine improvement in vision. Our results clearly demonstrated a marked improvement in the ability to detect and recognize letters following prolonged exposure to optical defocus. We ensured that ophthalmic change did not occur, and thus the phenomenon must be due to a neural compensation for the defocus condition. A second set of experiments measured contrast sensitivity and found a decrease in sensitivity to mid-range (5-25 cycles deg-1) spatial frequencies following exposure to optical defocus. The results of the two experiments may be explained by the unmasking of low contrast, high spatial frequency information via a two-stage process: (1) the pattern of relative channel outputs is maintained during optical defocus by the depression of mid-range spatial frequency channels; (2) channel outputs are pooled prior to the production of the final percept. The second set of experiments also provided some evidence of inter-ocular transfer, indicating that the adaptation process is occurring at binocular sites in the cortex.

Empirical and theoretical issues in the perception of time to contact.

Four questions concerning the perceptual source of information about time to contact (tc) are addressed: (a) What conditions are required for the optic variable tau to play a role in the perception of tc? (b) When these conditions are met, does tau alone provide sufficient information for accurate timing of interceptive actions? (c) Does a distance divided by velocity account of tc perception provide a convincing alternative to an account that is based on tau? (d) Is there any empirical evidence that distinguishes the two accounts? A "global" type of tau variable and a "local" type of tau variable are distinguished, each with different limitations. The discussion is largely concerned with local tau variables, 2 versions of which are identified. It is concluded that tau alone cannot provide sufficient information for skilled timing. An extended tau-based account presented in an earlier article (Tresilan, 1990) is discussed. It is argued that no extant empirical data can distinguish the extended account from the distance divided by velocity account.

Grip-load force coupling: a general control strategy for transporting objects.

The authors examined the coupling of grip force and load force during point-to-point and cyclic arm movements while holding an object in a variety of grips, including 1- and 2-handed grips and "inverted" grips. In all grips, grip force is modulated in phase with fluctuations in load force that are induced by the arm movement. The tight temporal coupling between grip force and load force seen when moving an object held in a precision grip (J. R. Flanagan, J. Tresilian, & A. M. Wing, 1993) is observed in other grips. The control of precision grip force during whole-body jumping movements was also investigated. Grip force was modulated in phase with changes in load force induced by jumping even though the arm's joint angles were fixed. The tight temporal coupling between grip force and load force during object transport reflects a general control strategy that is not specific to any particular grip or mode of transport. Models of the coordination of grasp and transport in prehensile behaviors are discussed.

Geometrical illusions can affect time-to-contact estimation and mimed prehension.

Results indicate that under some conditions the Sander parallelogram illusion can affect time-to-contact (TTC) estimation in a prediction-motion (PM) task and in an interceptive action (IA). The illusion also affected mimed manual prehension. The implication is that the timing of responses in the PM and IA tasks may involve an estimate of TTC that is based on the perceived dimensions of the environment. Further research is warranted in the development of models of perceived collision and of visually guided actions.

The preparation of reach to grasp movements in adults with Down syndrome.

The aim of this study was to determine the extent to which adults with Down syndrome (DS) are able to utilise advance information to prepare reach to grasp movements. The study comprised ten adults with DS; ten children matched to an individual in the group with DS on the basis of their intellectual ability, and twelve adult controls. The participants used their right hand to reach out and grasp illuminated perspex blocks. Four target blocks were positioned on a table surface, two to each side of the midsagittal plane. In the complete precue condition, participants were provided with information specifying the location of the target. In the partial precue condition, participants were given advance information indicating the location of the object relative to the midsagittal plane (left or right). In the null condition, advance information concerning the position of the target object was entirely ambiguous. It was found that both reaction times and movement times were greater for the participants with DS than for the adults without DS. The reaction times exhibited by individuals with DS in the complete precue condition were lower than those observed in the null condition, indicating that they had utilised advance information to prepare their movements. In the group with DS, when advance information specified only the location of the target object relative to the midline, reaction times were equivalent to those obtained when ambiguous information was given. In contrast, the adults without DS exhibited reaction times that were lower in both the complete and partial precue conditions when compared to the null condition. The pattern of results exhibited by the children was similar to that of the adults without DS. The movement times exhibited by all groups were not influenced by the precue condition. In summary, our findings indicate that individuals with DS are able to use advance information if it specifies precisely the location of the target object in order to prepare a reach to grasp movement. The group with DS were unable, however, to obtain the normal advantage of advance information specifying only one dimension of the movement goal (i.e., the position of an object relative to the body midline).

Manual interception of moving targets in two dimensions: performance and space-time accuracy.

We report results from four experiments that examined performance of an interceptive task that restricted movement of the hand and moving target to a horizontal plane. The task required accurate control over both where and when interception takes place. Three experiments studied the effects of four independent variables: target speed, target size, manipulandum size and movement amplitude. For small amplitude movements, small, fast targets were hit harder than larger slower ones and targets were hit harder with smaller manipulanda; movement time (MT) was unaffected by target size, but was shorter when the manipulandum was smaller. For larger amplitude movements, smaller, faster targets were also hit harder, but MTs tended to be greater when targets were smaller. The results support the idea that MT and peak movement speed can be independently controlled to some degree in order to meet the accuracy demands of the task. Analysis of the task showed that spatial and temporal accuracy demands are interdependent, indicating that the spatial and temporal variable errors should covary such that increases in one are accompanied by decreases in the other. This can be tested if there is no variation in interception location; which was not the case in the first three experiments. In a final experiment variation in interception location was restricted by requiring that the target be struck through an aperture. Both spatial and temporal variable errors could be estimated. As predicted, it was found that when spatial errors were small, temporal errors were large.

Systematic variation in performance of an interceptive action with changes in the temporal constraints.

People are highly skilled at intercepting moving objects and are capable of remarkably accurate timing. The timing accuracy required depends upon the period of time for which contact with a moving target is possible--the "time window" for successful interception. Studies of performance in an experimental interception task that allows this time window to be manipulated suggest that people change aspects of their performance (movement time, MT, and movement speed) in response to changes in the time window. However, this research did not establish whether the observed changes in performance were the results of a response to the time window per se or of independent responses to the quantities defining the time window (the size and speed of a moving target). Experiment 1 was designed to resolve this issue. The speed and size of the target were both varied, resulting in variations in the time window; MT was the primary dependent measure. Predictions of the hypothesis that people respond directly to changes in the time window were verified. Predictions of the alternative hypothesis that responses to changes in target speed and size are independent of one another were not supported. Experiment 2 examined how the type of performance change observed in Experiment 1 was affected by changing the time available for executing the interception. The time available and the target speed were varied, and MT was again the primary dependent measure. MT was smaller when there was less time available, and the effect of target speed (and hence the time window) on MT was also smaller, becoming undetectable at the shortest available time (0.4 s). The results of the two experiments are interpreted as providing information about the "rule" used to preprogramme movement parameters in anticipatory interceptive actions.

The preparation of reach-to-grasp movements in adults, children, and children with movement problems.

This study explored the use of advance information in the control of reach-to-grasp movements. The paradigm required participants to reach and grasp illuminated blocks with their right hand. Four target blocks were positioned on a table surface, two each side of the mid-saggital plane. In the complete precue condition, advance information precisely specified target location. In the partial precue condition, advance information indicated target location relative to the midsaggital plane (left or right). In the null condition, the advance information was entirely ambiguous. Participants produced fastest responses in the complete precue condition, intermediate response times in the partial condition, and the slowest responses in the null condition. This result was observed in adults and four groups of children including a group aged 4-6 years. In contrast, children with Developmental Coordination Disorder (DCD, n=11, aged 7--13 years) showed no advantage of partial precueing. Movement duration was determined by target location but was unaffected by precue condition. Movement duration was a clear function of age apart from children in the DCD group who showed equivalent movement times to those of the youngest children. These findings provide important insights into the control of reach-to-grasp movements and highlight that partial cues are exploited by children as young as 4 years but are not used in situations of abnormal development.

Perceptual and cognitive processes in time-to-contact estimation: analysis of prediction-motion and relative judgment tasks.

Three classes of task appear to involve time-to-contact (TTC) information: coincidence anticipation (CA) tasks, relative judgement (RJ) tasks, and interceptive actions (IAs). An important type of CA task used to study the perception of TTC is the prediction-motion (PM) task. The question of whether it is possible to study the perceptual processes involved in the timing of IAs using PM and RJ tasks is considered. A revised version of the tau hypothesis is proposed as an account of the perceptual information processing involved in the control of fast IAs. This draws on the distinction between "motor" and "cognitive" visual systems. It is argued that task variables affect whether "cognitive" information processing is involved in performance and can determine whether TTC information is used at all. Evidence is reviewed that suggests that PM and RJ tasks involve cognitive processing. It is argued that target viewing time, TTC at response initiation, amount of practice, and whether there is a period between target disappearance and response are task variables that determine whether cognitive processing will influence responding.

Four questions of time to contact: a critical examination of research on interceptive timing.

Four questions concerning the use and perception of time to contact, tc, are identified. (i) Is tc information used in the timing of interceptive actions? (ii) If so, what control strategies are used? (iii) What are the perceptual sources of tc information and which of them do people use? (iv) How is the information extracted by the perceptual systems? Research relevant to these questions is reviewed and analysed. In connection with question (i), theoretical work on the special case of catching a moving object is analysed. It is concluded that treatments of catching which involve the use of tc information provide the best account of timing. In connection with question (ii), two types of control strategy suggested in the literature are identified: an intermittent strategy and a continuous strategy. Evidence for a continuous strategy is reconsidered and shown to be at least as well if not better accounted for by an intermittent strategy. Other empirical evidence for intermittent control is also discussed. In connection with question (iii) a simple unifying method is outlined with which all tc information so far presented in the literature can be derived, and examples are given. The viability of various types of information as sources of tc is examined by considering the errors which would result from their use. Finally, in connection with question (iv) the role of 'looming detectors' in the extraction of tc information is considered. These are frequently proposed as mechanisms for extracting the tc information provided by Lee's optic variable, tau. The analysis provided indicates that, despite the existence of a well-known and popular theory, due mainly to Lee, about how interceptive actions are timed, very little is actually known about perceptual timing. It is not yet certain whether tc information is used in interceptive timing tasks, what kinds of control strategies are involved, what sources of information people use to time their actions, or what perceptual processing is involved in the extraction of tc information.

Correcting some misperceptions of time-to-collision: a critical note.

Stewart et al (1993, Perception 22 1227-1224) attempted to demonstrate that the ratio of angular speed to angular acceleration, [symbol see text]/[symbol see text], of the approaching object is preferred as the source of time-to-collision information to the ratio of visual angle to angular speed, [symbol see text]/[symbol see text]. It is demonstrated that this is not justified.

Analysis of recent empirical challenges to an account of interceptive timing.

How do we perceive how long it will be before we reach a certain place when running, driving, or skiing? How do we perceive how long it will be before a moving object reaches us or will arrive at a place where it can be hit or caught? These are questions of how we temporally coordinate our actions with a dynamic environment so as to control collision events. Much of the theoretical work on the control of these interceptive actions has been united in supposing that (1) timing is functionally separable from positioning and the two are controlled using different types of information; (2) timing is controlled using special-purpose time-to-arrival information; (3) the time-to-arrival information used for the timing of fast interceptive actions is a first-order approximation to the actual time-to-arrival, which does not take accelerations into account. Challenges to each of these suppositions have recently emerged, suggesting that a complete rethinking of how interceptions are controlled may be necessary. These challenges are analyzed in detail and it is shown that they are readily accommodated by a recent theory of interceptive timing based on the points just noted.

Retaining the equilibrium point hypothesis as an abstract description of the neuromuscular system.

The lambda version of the equilibrium point (EP) hypothesis for motor control is examined in light of recent criticisms of its various instantiations. Four important assumptions that have formed the basis for recent criticism are analyzed: First, the assumption that intact muscles possess invariant force-length characteristics (ICs). Second, that these ICs are of the same form in agonist-antagonist pairs. Third, that muscle control is monoparametric and that the control parameter, lambda, can be given a neurophysiological interpretation. Fourth, that reflex loop time delays and the known, asymmetric, nonlinear mechanical properties of muscles can be ignored. Mechanical and neurophysiological investigations of the neuromuscular system suggests that none of these assumptions is likely to be correct. This has been taken to mean that the EP hypothesis is oversimplified and a new approach is needed. It is argued that such an approach can be provided without rejecting the EP hypothesis, rather to regard it as an input-output description of muscle and associated segmental circuits. The operation of the segmental circuitry can be interpreted as having the function, at least in part, of compensating for a variety of nonlinearities and asymmetries such that the overall system implements the lambda-EP model equations.

Attention in action or obstruction of movement? A kinematic analysis of avoidance behavior in prehension.

Obstacle avoidance strategies are of two basic but interrelated types: moving around an obstacle to that body parts do not come too close, and slowing down. In reaching-to-grasp, avoidance may involve the transport component, the grasp formation component, or both. There has been little research that has directly examined obstacle avoidance strategies during reaches-to-grasp. Several recent reports describe experiments in which reaches-to-grasp were made when nontarget objects were present in the workspace. The effects of these nontargets were interpreted as being due to their distracting effects rather than their obstructing effects. The results of these studies are reinterpreted as being due to the non-target's obstructing effects. The obstacle interpretation is more parsimonious and better predicts the pattern of results than the distractor interpretation. Predictions of the obstacle interpretation were examined in an experiment in which participants were required to reach to grasp a target in the presence of another object in various locations. The results were exactly in line with the interpretation of the object as an obstacle and the data show how grasp and transport movements are subtly adjusted so as to avoid potential obstacles. It is proposed that people move so as not to bring body parts within a minimum preferred distance from nontarget objects within the workspace. What constitutes the preferred distance in a particular context appears to depend upon the speed of movement and a variety of psychological factors related to the cost that a person attaches to a collision.