The use of proprioception and tactile information in haptic search

To investigate how tactile and proprioceptive information are used in haptic object discrimination we conducted a haptic search task in which participants had to search for either a cylinder, a bar or a rotated cube within a grid of aligned cubes. Tactile information from one finger is enough to detect a cylinder amongst the cubes. For detecting a bar or a rotated cube amongst cubes touch alone is not enough. For the rotated cube this is evident because its shape is identical to that of the non-targets, so proprioception must provide information about the orientation of the fingers and hand when touching it. For the bar one either needs proprioceptive information about the distance and direction of a single finger’s movements along the surfaces, or proprioceptive information from several fingers when they touch it simultaneously. When using only one finger, search times for the bar were much longer than those for the other two targets. When the whole hand or both hands were used the search times were similar for all shapes. Most errors were made when searching for the rotated cube, probably due to systematic posture-related biases in judging orientation on the basis of proprioception. The results suggest that tactile and proprioceptive information are readily combined for shape discrimination.     

The Empathy Quotient (EQ) predicts perceived strength of bodily illusions and illusion-related sensations of pain

In this study we test the possible implications of high empathising skills on bodily self–other distinction by measuring the strength of a body ownership illusion and a related experience of illusory pain. One-hundred adult participants completed the Empathy Quotient (EQ) questionnaire. Twenty participants from the top quintile and 20 participants from the bottom quintile of the EQ distribution took part in a laboratory experiment. In the experiment, a classical Rubber Hand Illusion (RHI) was induced followed by the presentation of a series of painful stimuli to the rubber hand. Participants were asked to self-rate the strength of their subjective experience of the RHI and of the illusory pain. A proprioceptive location judgment on the position of the hidden hand was also required before and after RHI induction, to record drifts towards the rubber hand. We found a significant difference between high- and low-empathy participants in RHI and pain score. The EQ was not related with the proprioceptive location judgement drift. It thus appears to be a better predictor of subjective ownership feelings and phenomenological self–other merging than of the behavioural components of bodily illusions.     

Adaptation to visual rearrangement: Role of sensory discordance

The relative contributions of proprioceptive and efferent information in eliciting adaptation to visual rearrangement were studied under two conditions of visual stimulation. Subjects permitted sight of their forearm under normal room illumination showed significant adaptation when the forearm was (a) moved up and down under the action of tonic vibration reflexes, (b) voluntarily moved through the same trajectory at the same pace, (c) viewed while still, and (d) viewed while the margins of the elbow were vibrated. The reflex movement condition elicited significantly greater adaptation than the other conditions. Subjects allowed only sight of a point source of light attached to their hand showed significant adaptation when the forearm was (a) reflexly moved, (b) voluntarily moved through the same trajectory at the same rate, (c) passively moved, (d) still, and (e) vibrated while still. Less adaptation occurred as the amount of proprioceptive information about limb position was decreased. The adaptation elicited by voluntary movements of the forearm and by reflex movements did not differ significantly. It is concluded that corollary-discharge signals may not be crucial in adaptation to visual rearrangement; a more important factor appears to be discordance between proprioceptive and visual information.     

Prism adaptation in alternately exposed hands

We assessed intermanual transfer of the proprioceptive realignment aftereffects of prism adaptation in right-handers by examining alternate target pointing with the two hands for 40 successive trials, 20 with each hand. Adaptation for the right hand was not different as a function of exposure sequence order or postexposure test order, in contrast with adaptation for the left hand. Adaptation was greater for the left hand when the right hand started the alternate pointing than when the sequence of target-pointing movements started with the left hand. Also, the largest left-hand adaptation appeared when that hand was tested first after exposure. Terminal error during exposure varied in cycles for the two hands, converging on zero when the right hand led, but no difference appeared between the two hands when the left hand led. These results suggest that transfer of proprioceptive realignment occurs from the right to the left hand during both exposure and postexposure testing. Such transfer reflects the process of maintaining spatial alignment between the two hands. Normally, the left hand appears to be calibrated with the right-hand spatial map, and when the two hands are misaligned, the left-hand spatial map is realigned with the right-hand spatial map.     

Path information effects in visual and proprioceptive spatial learning

Objects in an environment are often encountered sequentially during spatial learning, forming a path along which object locations are experienced. The present study investigated the effect of spatial information conveyed through the path in visual and proprioceptive learning of a room-sized spatial layout, exploring whether different modalities differentially depend on the integrity of the path. Learning object locations along a coherent path was compared with learning them in a spatially random manner. Path integrity had little effect on visual learning, whereas learning with the coherent path produced better memory performance than random order learning for proprioceptive learning. These results suggest that path information has differential effects in visual and proprioceptive spatial learning, perhaps due to a difference in the way one establishes a reference frame for representing relative locations of objects.     

Effects on prism adaptation of duration and timing of visual feedback during pointing

In two experiments, we investigated the effects of duration of visual feedback of the pointing limb and the time (early to late) in the movement when the limb first becomes visible (timing of visual feedback). Timing, rather than duration of visual feedback, proved to have the greater effect on the relative magnitude of visual and proprioceptive adaptation. Visual adaptation increased smoothly with feedback delay, but corresponding decreases in proprioceptive adaptation underwent an additional sharp change when feedback was delayed until about three-fourths of the way to the terminal limb position. These results are consistent with the idea that visual and proprioceptive adaptation are mediated by exclusive processes. Change in the limb position sense (i.e., proprioceptive adaptation) may be produced by visual guidance of the pointing limb, and view of the limb early in the pointing movement seems to be critical for such visual guidance. The limb may be ballistically released as it nears the terminal position, and, thereafter, any opportunity for visual guidance (i.e., view of the limb) is not effective. On the other hand, change in the eye position sense (i.e., visual adaptation) may be mediated by proprioceptive guidance of the eye; the eyes may track the imaged position of the nonvisible limb. Such proprioceptive guidance seems to be solely a function of the distance moved before the limb becomes visible.     

Active and passive-touch during interpersonal multisensory stimulation change self–other boundaries

In the “enfacement” illusion seeing an unfamiliar face being touched at the same time as one’s own face evokes changes in self-face recognition. We investigated the contribution of proprioceptive and motor signals derived from self-generated actions in the sensory-driven malleability of self–other boundaries during the “enfacement” illusion. Changes in self-face recognition during active- and passive-touch interpersonal visuo-tactile stimulation were quantified by means of psychophysical and psychometric tasks. Active- and passive-touch evoked comparable changes in the categorical boundaries of self–other distinction, changing the extent to which the other is assimilated into the mental self-representation. Actively touching or simply feeling touch on one’s own face with concurrent observed touch on someone else’s face seems to elicit comparable changes in self-recognition, suggesting that afferent input might be sufficient for updating one’s body-image, although some components of the experience of self-identification seem to be more affected by passive- than by active-touch.     

Proprioception as a Basis for the Temporal Anticipation of Motor Responses,

The present study attempted to distinguish between the proprioceptive Decay and proprioceptive Input Hypotheses of motor timing behavior. The hypothesis that an increased level of proprioceptive cues results in more proficient temporal anticipation was also tested. 90 male Ss were required to anticipate (no preview) the coincidence of a moving pointer and a stationary one. The timing response was executed with the right hand while 3 levels of proprioceptive cues were indirectly manipulated in the left arm. While results were unable to distinguish between the hypotheses, support was found for the hypothesis that an increased level of proprioceptive feedback administered during the interval can increase anticipatory response consistency.     

Additivity of components of prismatic adaptation

Ss pointed with each hand at a light or at the unseen toe and looked in the direction of the unseen toe before, during, and after training one arm to point to a visual target which was progressively displaced to one side by a prism. Results show that a proprioceptive change in the trained arm is a universal component of the adaptation. When a change in the eye-head system occurs, it and the proprioceptive change in the arm sum to the total adaptation and it is accompanied by a predictable degree of intermanual transfer of the adaptation, as a felt-position theory of adaptation would predict. However, when there is no change in the eye-head system, the proprioceptive shift is not always sufficient to account for the total adaptive shift.     

The role of proprioception in the perception and control of human movement: Toward a theoretical reassessment

The theoretical role of proprioception in the perception and control of human movement is elusive because of the obvious inability to manipulate experimentally the various receptive systems. Individuals who have had the metacarpophalangeal joint and joint capsule removed and replaced with silastic inserts afford a unique opportunity to evaluate a principal source of proprioception, namely, slowly adapting joint afferents. In a set of experiments, we show that such individuals show no deficits in finger localization following joint replacement. We take this and other complementary findings as a preliminary basis for proposing a dynamic rather than kinematic account of movement production. In addition, we provide a reconceptualization of the function of proprioceptive information in the CNS. Our arguments focus on proprioceptive inputs as tuning or modulating interneuronal pools rather than providing dimension-pecific information to the brain, as is commonly assumed.     

Contingent orienting or contingent capture: A size singleton matching the target–distractor size relation cannot capture attention

In the present study, we examined whether a peripheral size-singleton distractor that matches the target–distractor size relation can capture attention and disrupt central target identification. Three experiments consistently showed that a size singleton that matches the target–distractor size relation cannot capture attention when it appears outside of the attentional window, even though the same size singleton produces a cuing effect. In addition, a color singleton that matches the target color, instead of a size singleton that matches the target–distractor size relation, captures attention when it is outside of the attentional window. Thus, a size-relation-matched distractor is much weaker than a color-matched distractor in capturing attention and cannot capture attention when the distractor appears outside of the attentional window.     

Effects on Prism Adaptation of Duration and Timing of Visual Feedback During Pointing

In two experiments, we investigated the effects of duration of visual feedback of the pointing limb and the time (early to late) in the movement when the limb first becomes visible (timing of visual feedback). Timing, rather than duration of visual feedback, proved to have the greater effect on the relative magnitude of visual and proprioceptive adaptation. Visual adaptation increased smoothly with feedback delay, but corresponding decreases in proprioceptive adaptation underwent an additional sharp change when feedback was delayed until about three-fourths of the way to the terminal limb position. These results are consistent with the idea that visual and proprioceptive adaptation are mediated by exclusive processes. Change in the limb position sense (i.e., proprioceptive adaptation) may be produced by visual guidance of the pointing limb, and view of the limb early in the pointing movement seems to be critical for such visual guidance. The limb may be ballistically “released“ as it nears the terminal position, and, thereafter, any opportunity for visual guidance (i.e., view of the limb) is not effective. On the other hand, change in the eye position sense (i.e., visual adaptation) may be mediated by proprioceptive guidance of the eye; the eyes may track the imaged position of the nonvisible limb. Such proprioceptive guidance seems to be solely a function of the distance moved before the limb becomes visible.     

Proprioception plays a different role for sensorimotor adaptation to different distortions

If proprioceptive feedback is degraded by agonist-antagonist muscle vibration, then adaptation to rotated vision remains intact while adaptation to a velocity-dependent force field worsens. Here we evaluate whether this differential effect of vibration is related to the physical nature of the distortion - visual versus mechanical - or to their kinematic coupling to the subjects’ hand - velocity versus position dependent. Subjects adapted to a velocity-dependent visual distortion, to a position-dependent force, or to a velocity-dependent force; one half of the subjects adapted with, and the other half without agonist-antagonist vibration at the wrist, elbow, and shoulder. We found, as before, that vibration slowed down adaptation to a velocity-dependent force. However, vibration did not modify adaptation to the other two distortions, nor did it influence the aftereffects of any distortion. From this we conclude that intact proprioception supports strategic compensatory processes when proprioceptive signals agree with visual ones, and provide relevant (dynamic) information not available to the visual system.     

Effects of Pointing Rate and Availability of Visual Feedback on Visual and Proprioceptive Components of Prism Adaptation

While looking through laterally displacing prisms, subjects pointed 60 times straight ahead of their nose at a rate of one complete movement every 2 or 3 s, with visual feedback available early in the pointing movement or delayed until the end of the movement. Sagittal pointing was paced such that movement speed covaried with pointing rate. Aftereffect measures (obtained after every 10 pointing trials) showed that when the limb became visible early in a pointing movement, proprioceptive adaptation was greater than visual, but when visual feedback was delayed until the end of the movement, the reverse was true. This effect occurred only with the 3-s pointing rate, however. With the 2-s pointing rate, adaptation was predominately proprioceptive in nature, regardless of feedback availability. Independent of the availability of visual feedback, visual adaptation developed more quickly with 3-s pointing, whereas proprioceptive adaptation developed more rapidly with 2-s pointing. These results are discussed in terms of a model of perceptual-motor organization in which the direction of coordinative (guidance) linkage between eye-head (visual) and hand-head (proprioceptive) systems (and consequently the locus of discordance registration and adaptive recalibration) is determined jointly by pointing rate and feedback availability. An additional effect of pointing rate is to determine the rate of discordant inputs. Maximal adaptive recalibration occurs when the input (pointing) rate matches the time constant of the adaptive encoder in the guided system.     

Temporal Distribution of Attention During a Throwing Motion

A reaction-time measure of attention was employed during five segments of a simultaneous dart throw: premovement decision, immediately after recoil was begun, maximum recoil, immediately after forward movement was begun, and after release of the dart. Reaction time was significantly longer when the stimulus was presented during the first or second segment than during later segments. Reaction time during premovement was not consistently greater than that immediately after movement was begun, indicating that information is being sampled at a high rate after movement commences. It appears that monitoring of proprioceptive feedback after movement has begun, and initiation of corrective submovements, is necessary for skillful performance of dart throwing.     

Eye movement control during reading: Evidence for direct control

In three experiments, subjects read text as their eye movements were monitored and display changes in the text were made contingent upon the eye movements. In one experiment, a window of text moved in synchrony with the eyes. In one condition, the size of the window was constant from fixation to fixation, while in the other condition the size of the window varied from fixation to fixation. In the other experiments, a visual mask was presented at the end of each saccade which delayed the onset of the text, and the length of the delay was varied. The pattern of eye movements was influenced by both the size of the window and the delay of the onset of the text, even when the window size or text delay was varying from fixation to fixation. However, there was also evidence that saccade length was affected by the size of the window on the prior fixation and that certain decisions to move the eye are programmed either before the fixation begins or are programmed during the fixation but without regard to the text fixated on. The results thus provide support for a mixed control model of eye movements in reading, in which decisions about when and where to move the eyes are based on information from the current fixation, the prior fixations, and possibly, other sources as well.     

A cross-modal interference effect in grasping objects

The aim of the present paper was to investigate how the kinematics of a hand reaching toward a visual target would be influenced by haptic and proprioceptive input from an unseen distractor actively grasped in the other, nonreaching hand. The main results were that the amplitude of maximum grip aperture was smaller and the time to maximum grip aperture was earlier when the distractor was smaller than the target. The interference effect from the distractor was similar for both hands as they reached. Furthermore, results from a vibrating-distractor condition for passive tactile input revealed that the interference effects were evident only when the distractor was actively grasped. We suggest that neural processing of proprioceptive and tactile information relevant to distractor size produced the observed interference effects. We also emphasize the importance of active manipulation of the distractor stimulus in eliciting such interference effects.     

The case for proprioception

In formulating a theory of perception that does justice to the embodied and enactive nature of perceptual experience, proprioception can play a valuable role. Since proprioception is necessarily embodied, and since proprioceptive experience is particularly integrated with one’s bodily actions, it seems clear that proprioception, in addition to, e.g., vision or audition, can provide us with valuable insights into the role of an agent’s corporal skills and capacities in constituting or structuring perceptual experience. However, if we are going to have the opportunity to argue from analogy with proprioception to vision, audition, touch, taste, or smell, then it is necessary to eschew any doubts about the legitimacy of proprioception’s inclusion into the category of perceptual modalities. To this end, in this article, I (1) respond to two arguments that Shaun Gallagher (2003) presents in “Bodily self-awareness and objectperception” against proprioception’s ability to meet the criteria of object perception, (2) present a diagnosis of Gallagher’s position by locating a misunderstanding in the distinction between proprioceptive information and proprioceptive awareness, and (3) show that treating proprioception as a perceptual modality allows us to account for the interaction of proprioception with the other sensory modalities, to apply the lessons we learn from proprioception to the other sensory modalities, and to account for proprioceptive learning. Finally, (4) I examine Sydney Shoemaker’s (1994) identification constraint and suggest that a full-fledged notion of object-hood is unnecessary to ground a theory of perception.     

Central fatigue mechanisms are responsible for decreases in hand proprioceptive acuity following shoulder muscle fatigue

Muscle fatigue is a complex phenomenon, consisting of central and peripheral mechanisms which contribute to local and systemic changes in motor performance. In particular, it has been demonstrated that afferent processing in the fatigued muscle (e.g., shoulder), as well as in surrounding or distal muscles (e.g., hand) can be altered by fatigue. Currently, it is unclear how proximal muscle fatigue affects proprioceptive acuity of the distal limb. The purpose of the present study was to assess the effects of shoulder muscle fatigue on participants’ ability to judge the location of their hand using only proprioceptive cues. Participants’ (N = 16) limbs were moved outwards by a robot manipulandum and they were instructed to estimate the position of their hand relative to one of four visual reference targets (two near, two far). This estimation task was completed before and after a repetitive pointing task was performed to fatigue the shoulder muscles. To assess central versus peripheral effects of fatigue on the distal limb, the right shoulder was fatigued and proprioceptive acuity of the left and right hands were tested. Results showed that there was a significant decrease in the accuracy of proprioceptive estimates for both hands after the right shoulder was fatigued, with no change in the precision of proprioceptive estimates. A control experiment (N = 8), in which participants completed the proprioceptive estimation task before and after a period of quiet sitting, ruled out the possibility that the bilateral changes in proprioceptive accuracy were due to a practice effect. Together, these results indicate that shoulder muscle fatigue decreases proprioceptive acuity in both hands, suggesting that central fatigue mechanisms are primarily responsible for changes in afferent feedback processing of the distal upper limb.     

Novel metaphors appear anomalous at least momentarily: evidence from N400

This study addresses a central question in perception of novel figurative language: whether it is interpreted intelligently and figuratively immediately, or only after a literal interpretation fails. Eighty sentence frames that could plausibly end with a literal, truly anomalous, or figurative word were created. After validation for meaningfulness and figurativeness, the 240 sentences were presented to 11 subjects for event related potential (ERP) recording. ERP's first 200 ms is believed to reflect the structuring of the input; the prominence of a dip at around 400 ms (N400) is said to relate inversely to how expected a word is. Results showed no difference between anomalous and metaphoric ERPs in the early window, metaphoric and literal ERPs converging 300-500 ms after the ending, and significant N400s only for anomalous endings. A follow-up study showed that the metaphoric endings were less frequent (in standardized word norms) than were the anomalous and literal endings and that there were significant differences in cloze probabilities (determined from 24 new subjects) among the three ending types: literal > metaphoric > anomalous. It is possible that the low frequency of the metaphoric element and lower cloze probability of the anomalous one contributed to the processes reflected in the early window, while the incongruity and near-zero cloze probability of the anomalous endings produced an N400 effect in them alone. The structure or parse derived for metaphor during the early window appears to yield a preliminary interpretation suggesting anomaly, while semantic analysis reflected in the later window renders a plausible figurative interpretation.