Adaptation to altered visual—vestibular feedback: Mechanisms of maintenance and recovery

Adaptation of perceived movement during head motion (apparent concomitant motion, ACM) and the subsequent elimination of adaptation were studied in two experiments. During the adaptation phase of both experiments, subjects performed voluntary 1-Hz head oscillations for 6 min while fixating a stimulus moving either in the same (with) direction as or the opposite (against) direction of head movements. In Experiment 1, ACM adaptation was measured following either a 1- or a 4-min delay after the adaptation phase. Results indicated some loss of adaptation during the additional 3-min delay, demonstrating a tendency of the system linking head and image to return to its preadaptation state following removal of an adaptation stimulus. In Experiment 2, subjects viewed a stimulus after adaptation that appeared to move minimally in the same manner as the adaptation stimulus during 3 min of head oscillations. No loss of adaptation was measured in these subjects between the beginning and the end of the 3-min interval. In another condition, subjects viewed a stimulus that appeared to move alternately in the same direction as and in the opposite direction of the adaptation stimulus during a similar 3-min interval following adaptation. ACM adaptation was substantially reduced during this 3-min interval. These results implicate two mechanisms that operate to either maintain or eliminate ACM adaptation. One is passive and operates in the absence of visual feedback to eliminate the short-term adapted state, and the other responds to postadaptation visual feedback.     

On counteradaptation

Often adaptation to artificially altered stimulation takes place because veridical stimulation that produces the same perceptual property that is produced by the altered stimulation is also received. In these cases, an assimilation of the two perceptual processes produced by the two different stimulations (the altered and the veridical) is supposed to be responsible for the adaptation that is achieved. This hypothesis, which was formulated by Wallach and Karsh (1963), would be confirmed by demonstrating a modification of the perceptual process produced by veridical stimulation rather than the one produced by the altered stimulation. We demonstrated this by having S observe in the dark for 20 min a luminous figure that objectively expanded as it moved toward S and contracted as it moved away. But instead of testing for changes in size perception as such, we tested for a change in the relation between accommodation and convergence on the one hand and registered distance on the other. In one experiment, such a change was measured by obtaining estimates of perceived size and depth before and after the adaptation period. Highly significant changes of size and significantly greater changes of stereoscopic depth were obtained. Inasmuch as stereoscopic vision was totally absent from the adaptation conditions, the change in stereoscopic depth that was larger than the size change can only be ascribed to a change in registered distance. In another experiment, we tested for a change in distance by having S point from the side to a vertical line, before and again after the adaptation period, under conditions where only accommodation and convergence could serve as distance cues. Significant changes in the pointing distance were measured, indicating more directly a change in the relation between these oculomotor adjustments and perceived distance. We propose the term counteradaptation for such modification of a perceptual process away from veridicality.     

Adaptation in distance perception based on oculomotor cues

Accommodation and convergence primarily serve to adjust the eyes to the distance of the object viewed, but, once made, these oculomotor adjustments serve as cues for the object’s distance. Experiments are reported that show that the relation between oculomotor adjustments and the distances they signify can be changed by adaptation to glasses that cause alteration in the oculomotor adjustments with which objects are viewed. This changed relation manifested itself in marked alterations of size perception. Wearing, for 30 min, glasses that caused a change in accommodation and convergence corresponding to a smaller object distance and equivalent to 1.5 lens diopters caused subsequent mean size increases that ranged from 50% to 65%. Adaptation to glasses that changed oculomotor adjustments in the same amount but in the opposite direction resulted in decreases in perceived sizes that varied from 18% to 40%, dependent on the distance of the test object. These were the results of size estimates obtained before and after the adaptation period under conditions where only accommodation and convergence served as cues for distance. A newly developed test of size perception was also used, in which S adjusted the size of the projected image of an array of familiar objects on a screen until the size of the objects appeared normal. Again, such adjustments were made before and after the adaptation period, and size differences were obtained that were in the direction to be expected of adaptation and varied in amount between 12% and 33%, dependent on the distance of the screen. The reason for the different amounts of size change measured by the two kinds of tests was investigated.     

Oculomotor adjustments in darkness and the specific distance tendency

Two experiments were performed to investigate the relationship between the oculomotor adjustments assumed in total darkness and perceived distance under reduced visual conditions. Experiment I compared the dark focus of accommodation with the perceived distance of a monocular light point presented in a dark environment. Experiment II compared the convergence angle assumed in darkness (dark convergence) with the perceived distance of the light point. Both accommodation and convergence were found to assume intermediate values in darkness. Perceived distance of the monocular light point was significantly correlated with dark convergence and unrelated to the dark focus of accommodation. It was suggested that ocular vergence is a major determinant of perceived distance under reduced visual conditions, and thus provides a possible mechanism for the specific distance tendency.     

A comparison of accommodative and fusional convergence as cues to distance

The effectiveness of fusional as compared with accommodative convergence (with accommodation present in both cases) in determining perceived distance was investigated in this study. Luminous frames of two different visual angles at a nearly constant distance were viewed binocularly to provide fusional convergence and monocularly to provide accommodative convergence. Although some differences in reported size and distance of the frames occurred on the first presentations for binocular as compared to monocular observation, the most systematic differences between these two types of observation were present for the second (successive) presentations of the two frame sizes to the same Os. This is attributed to the relative size cue to distance occurring as a function of the different retinal sizes on the successive presentations. It was found that this relative size cue was more effective in modifying the perceived size and distance of the second presentations for monocular than for binocular observation. It is suggested that this reflects the greater effectiveness as a cue to distance of fusional as compared with accommodative convergence. This conclusion is of importance for studies concerned with the evaluation of convergence as a determiner of perceived distance.     

Oculomotor adjustments and size-distance perception

The relationship between perceived size and distance and oculomotor adjustments were assessed in two experiments. In both experiments, Ss were required to make scalar linear size, angular size, and distance judgments of stimuli subtending a constant retinal image size at different levels of convergence. The results of the first experiment indicate that the perceived linear size, angular size, and distance of the stimulus decreased with increased convergence, the decrease in perceived linear size being greater than that of perceived angular size. While again showing a decrease in perceived linear and angular size, the results of the second experiment also show that there was a smaller decrease in perceived distance with increased convergence when Ss continued to view the stimulus as convergence was changed than when they did not view the stimulus as convergence was changed. The implications these results have for size and distance perception are discussed.     

The nature of adaptation in distance perception based on oculomotor cues

In previous work by the senior authors, brief adaptation to glasses that changed the accommodation and convergence with which objects were seen resulted in large alterations in size perception. Here, two further effects of such adaptation are reported: alterations in stereoscopic depth perception and a change when distance is represented by a response of S’s arm. We believe that the three effects are manifestations of one primary effect, an alteration of the relation between accommodation and convergence on the one hand and the distance they represent in the nervous system (registered distance) on the other. This view was supported by the results of two experiments, each of which demonstrated that the alterations in stereoscopic depth perception could be obtained after adaptation periods which had provided no opportunity to use stereoscopic vision, and that the adaptation effect was larger for depth perception than for size perception when it was obtained under the same conditions; the latter finding was expected if both effects resulted from the same change in registered distance. In three of the five experiments here reported, the variety of cues that could represent veridical distance during the adaptation period was limited. In one condition of adaptation, only the pattern of growth of the retinal images of objects that S approached and the kinesthetic cues for S’s locomotion served as cues to veridical distance. In two other conditions S remained immobile. In one of these, only the perspective distortion in the projection of the scene that S viewed mediated veridical distance, and in the other one familiar objects of normal size were successively illuminated in an otherwise totally dark field, conditions from which opportunities to use stereoscopic vision were again absent. After exposure to each of these adaptation conditions, adaptive changes in perceived size and larger ones in perceived stereoscopic depth were obtained. Because we found that familiar size may serve as the sole indicator of veridical distance in an adaptation process, we concluded that it can function as a perceptual as distinguished from an inferential cue to distance.     

Visual angle as a determinant of perceived interobject distance

Two new experiments and a reanalysis of Toye’s (1986)data are used to examine the relationship between true distance and perceived distance in natural scenes. In the first experiment, 8 subjects estimated 78 interobject distances, formed by all pairs of 13 objects, while viewing the objects from a fixed position. The results showed that estimated distance is a linear function of the visual angle between objects as well as of the true distance. This relationship results in distances perpendicular to the line of sight being overestimated in relation to true distances and to distances parallel to the line of sight. These findings were confirmed by reanalysis of a comparable data set from Toye. Since changes in the visual angle can come about through changes in alignment with the line of sight, viewing distance, or interobject distance, Experiment 2 was designed to determine whether the visual angle effect was due to one of these, or whether it was an independent effect. In Experiment 2, 8 subjects estimated six interobject distances from 12 viewing positions. The results showed that visual angle predicted estimated distance independently of how the change in visual angle came about, suggesting that the greater the visual angle between objects, the more their separation is overestimated.     

Distance adaptation depends upon plasticity in the oculomotor control system

Maintaining binocular fixation on a target at 20 cm in the absence of secondary cues to distance produced changes in apparent distance and lateral phoria. Positive lenses of 0, .5, 2.0, 3.5, and 5.0 spherical diopters (SD) were used to manipulate the level of accommodative convergence in force during the period of maintained fixation. An inverse relationship was found between the stimulus to accommodation and the magnitude of the induced esophoria, the phoria being linearly related to an increase in apparent distance. The distance aftereffect obtained in the condition with the lowest net accommodative stimulus li.e., 0 D) equaled that typically produced by base-out prism adaptation with full secondary cues to distance available. In a second experiment, subjects walked through a well-lit hallway while viewing through a pair of 5h base-out prisms. It was shown that increasing the stimulus to accommodation by adding negative lenses of 0, 1.5, 3.5, and 5.5 SD reduced the adaptive change in apparent distance, as well as the change in phoria produced by the conventional base-out prism adaptation paradigm. It was concluded that a change in the resting tonus of the disparity vergence system underlies such adaptation, rather than recalibration of the oculomotor cues to distance. Monocular exposure data indicated that a small change in the tonus control for the accommodative system may be present as well.     

Oculomotor adjustments and size constancy

The effect on matched size of the oculomotor adjustments was determined by stimulation and relaxation of accommodation and convergence by means of spherical lenses. The normal coupling between accommodation and convergence was maintained by introducing the amount of convergence appropriate to the lens power and each S’s interpupillary distance. Data indicate that the oculomotor adjustments are adequate to account for size constancy up to approximately 1 m, beyond which their effect progressively decreases. The actual accommodation in force was assessed by means of the laser scintillation technique. It was determined that the magnitude of accommodation responds accurately to the spherical lens introduced up to about 1 m observation distance, beyond which underaccommodation was noted. Examination of the matched size as a function of the actual accommodation distance reveals a very close correspondence to the size constancy prediction up to about 1 m.     

Spiral aftereffects after long-term delay

Tested the effects of three kinds of postadapation stimuli on short- and long-term spiral aftereffects. Magnitude and duration of short-term aftereffects were significantly reduced by 5 min. of no visual stimulation preceding inspection of the postadaptation field. Results do not support previous findings of long-term aftereffects.     

Effect of disparity and viewing distance on perceived depth

It is shown that veridical depth perception presupposes the processing of both the magnitude of retinal disparity and observation distance according to a square-law function specified by the underlying geometrical stimulus relations. In the present study, after testing its existence, this constancy of depth perception was investigated by measuring perceived depth as a function of retinal disparity and observation distance. In addition, the relative effectiveness of convergence and accommodation as possible indicators of distance was examined through a conflicting-cues paradigm. It was shown that in the perception of depth the visual system computes distance by taking into account the convergence parameter only, rather than that of accommodation or of both.     

Target distance and adaptation in distance perception in the constancy of visual direction

Hay and Sawyer recently demonstrated that the constancy of visual direction (CVD) also operates for near targets. A luminous spot in the dark, 40 cm from the eyes, was perceived as stationary when S nodded his head. This implies that CVD takes target distance, as well as head rotation, into account as a stationary environment is perceived during head movements. Distance is a variable in CVD because, during a turning or nodding of the head, the eyes become displaced relative to the main target direction, the line between the target and the rotation axis of the head. This displacement of the eyes during head rotation causes an additional change in the target direction, i.e., a total angular change greater than the angle of the head rotation. The extent of this additional angular displacement is greater the nearer the target. We demonstrated that the natural combination of accommodation and convergence can supply the information needed by the nervous system to compensate for this additional target displacement. We also found that wearing glasses that alter the relation between these oculomotor adjustments and target distance produces an adaptation in CVD. An adaptation period of 1.5 h produced a large adaptation effect. This effect was not entirely accounted for by an adaptation in distance perception. Measurements of the alteration between oculomotor cues and registered distance with two kinds of tests for distance perception yielded effects significantly smaller than the effect measured with the CVD test. We concluded that the wearing of the glasses had also produced an adaptation within CVD.     

Adaptation to different mouth shapes influences visual perception of ambiguous lip speech

We investigated the effects of adaptation to mouth shapes associated with different spoken sounds (sustained /m/ or /u/) on visual perception of lip speech. Participants were significantly more likely to label ambiguous faces on an /m/-to-/u/ continuum as saying /u/ following adaptation to /m/ mouth shapes than they were in a preadaptation test. By contrast, participants were significantly less likely to label the ambiguous faces as saying /u/ following adaptation to /u/ mouth shapes than they were in a preadaptation test. The magnitude of these aftereffects was equivalent when the same individual was shown in the adaptation and test phases of the experiment and when different individuals were presented in the adaptation and test phases. These findings present novel evidence that adaptation to natural variations in facial appearance influences face perception, and they extend previous research on face aftereffects to visual perception of lip speech.     

Modification of depth and distance perception caused by long-term wearing of left-right reversing spectacles

For 35 to 39 days, four observers wore continuously left-right reversing spectacles which pseudoscopically reverse the order of binocular disparity and direction of convergence. In three tests, we investigated how the visual system copes with the transformation of depth and distance information due to the reversing spectacles. In stereogram observation, after a few days of wearing the spectacles. the observers sometimes perceived a depth order which was opposite to the depth order that they had perceived in the pre-spectacle-wearing period. Monocular depth cues contributed more to depth perception in the spectacle-wearing period than they did in the pre-spectacle-wearing period. While the perceived distance significantly decreased during the spectacle-wearing period, we found no evidence of adaptive change in distance perception. The results indicate that the visual system adapts itself to the transformed situation by not only changing the processing of disparity but also by changing the relative efficiency of each cue in determining apparent depth.     

Understanding the structural determinants of object confusion in memory: an assessment of psychophysical approaches to estimating visual similarity

Computer-generated shapes varying on visual dimensions such as curvature, tapering, and thickness have been used to investigate identification deficits in the category-specific visual agnosia (CSVA) Patient E.L.M.. However, whether the implemented variations on each of these dimensions were perceived by novice observers as "similar amounts of change" is unknown. To estimate distance in psychophysical shape space, sets of shapes were developed using two different scaling methods--an objective method based on visual search, and a subjective method based on judgments of similarity--and a third approach that did not involve scaling. How well each method estimated psychophysical shape space was assessed by measuring the confusions within memory among the shapes. The results suggested that, although neither of the approaches perfectly reflected psychophysical shape space, subjective scaling was a better estimator of distance in psychophysical shape space than were other approaches. The number of confusions produced on each set of shapes was used to develop a new set of shapes that accurately estimated distance in psychophysical shape space. These results suggest that a combination of approaches is preferable in order to accurately estimate distance in psychophysical shape space.     

Binocular cues in the perception of distance of a point source of light

The ability to make egocentric distance estimates of a single point source of light, seen in darkness and without the cues of changing size and luminance, was investigated in sixteen observers. The attenuation required to maintain constant luminance, when the target was viewed from different distances, was shown to follow the inverse square law providing the angle subtended by the light was less than 20 s arc. Distance changes were also simulated by means of a split mirror which produced vergence cues, or by test lenses to provide accommodation cues. Over the range 0.5 to 9.2 m distance estimates were surprisingly accurate, although there was some overestimation of near and underestimation of far distances. Most observers made good judgements when only convergence cues were varied, whereas no observers made consistently good judgements when only accommodation cues were varied. The difficulties are discussed in terms of the accommodation-convergence link. When distance was simulated by changing convergence and accommodation cues, estimates were not as good as when real distance was changed. Since good estimates were made with brief target exposures, these judgements were not based on subsequent convergence or accommodation changes. It is suggested that the metric or reference against which the apparently absolute judgements were made was the efferent demand signal associated with a 'resting' position of convergence in darkness.     

Preferred gaze direction and the vergence system at rest]

In two experiments the effects of binocular vergence, head tilt, and prolonged gaze inclination on the preferred vertical gaze direction were studied. Concurrently, dark vergence was measured to test a hypothesis on the effect that binocular vergence has on the preferred vertical direction of gaze. The previously reported influence of observation distance on the preferred gaze inclination was replicated. When the head was tilted backward, the preferred gaze inclination was lower and the effect of observation distance was reduced; when the head was tilted forward, the preferred gaze inclination was higher and the effect of observation distance was increased. After prolonged upward or downward gaze inclination the preferred vertical gaze direction was shifted up or down, respectively, while the effect of observation distance remained constant. Overall the change of the preferred vertical direction of gaze upon binocular fixation of near stimuli was accompanied by a reduced discrepancy between convergence and the resting state of the vergence system; additional implications of the hypothesis on the relation between individual resting vergence and the effect of observation distance on the preferred inclination of gaze could not be substantiated.     

Perceived size and perceived distance in stereoscopic vision and an analysis of their causal relations

Effects of visual angle and convergence upon the perceived sizes and perceived distances of a familiar object (playing card) and a nonrepresentational object (blank white card) were investigated by means of a projector stereoscope with polarizing filters. The results obtained with six Ss indicated that size estimates increased nearly proportionally as the visual angle increased and decreased nearly linearly as the convergence increased. Distance estimates decreased nearly linearly as either the visual angle or the convergence increased. The ratio of the size estimate to the distance estimate for a given visual angle was almost constant irrespective of convergence. In this sense, the size-distance invariance hypothesis held. No clear effect of familiarity was found. Partial correlations were used to discriminate direct and indirect causal relationships between the stimulus variables and perceptual estimates. Both perceived size and perceived distance were found to be determined directly by the two stimulus variables, but to be mutually related only indirectly.     

Visual and proprioceptive adaptation to altered oculomotor adjustments

Adaptation to spectacles that alter in equivalent amounts the accommodation and the convergence with which objects are viewed was produced under two conditions. In one, S alternately pushed away or pulled toward him a screen that exhibited only a single vertical contour while wearing glaaaes that caused decreases in accommodation and convergence equivalent to 1.5 lens diopters. Here kinesthesis for these arm movements provided the only veridical distance cues, A small, but highly significant, adaptation effect was obtained with a teat in which S, before and after the adaptation period, pointed to the location of a test line in the distance dimension. Corresponding tests consisting in size and in depth estimates did not show an adaptation effect. In the other condition of adaptation, S moved objects by hand toward and away from himself while wearing spectacles that increased accommodation and convergence by the equivalent of 1.5 lens diopters. In addition to the altered oculomotor cues, some veridical visual cues for distance such as are caused by perspective were present. This condition yielded changes in size and depth estimates indicative of an adaptation in visual distance perception and a larger effect of adaptation measured by the pointing test. We concluded that the excess of the adaptation effect measured by pointing over that measured by size estimation represents an adaptation in proprioception, as did the pointing effect produced by our first adaptation condition.