Copyright 2001 by the American Psychological Association, Inc. 0096-1523/01/S5.00 DOI: 10.1037//O096-1523.27.1.206 Journal of Experimental Psychology: Human Perception and Performance 2001, Vol. 27, No. 1, 206-217 The Tilt-Constancy Theory of Visual Illusions William Prinzmetal Diane M. Beck University of California, Berkeley University College London The authors argue that changes in the perception of vertical and horizontal caused by local visual cues can account for many classical visual illusions. Because the perception of orientation is influenced more by visual cues than gravity-based cues when the observer is tilted (e.g., S. E. Asch & H. A. Witkin, 1948), the authors predicted that the strength of many visual illusions would increase when observers were tilted 30°. The magnitude of Zollner, Poggendorff, and Ponzo illusions and the tilt-induction effect substantially increased when observers were tilted. In contrast, the MiiUer-Lyer illusion and a size constancy illusion, which are not related to orientation perception, were not affected by body orientation. Other theoretical approaches do not predict the obtained pattern of results. shifts in gravity in relation to body position (Howard, 1998; Howard & Childerson, 1994), we call our new account of visual illusions the tilt-constancy theory. We first illustrate how local visual cues to orientation can account for many illusions. Second, we review the literature relevant to our claim. Finally, we present evidence in support of the tilt-constancy theory that cannot be explained by other theories. The effect of visual context on orientation perception was dramatically illustrated in classic studies by Asch and Witkin (1948). Observers viewed a room that was tilted 22° in the picture plane and were asked to adjust a rod so that it was oriented horizontally or vertically with respect to gravity. The average setting was distorted 15° in the direction of the tilt of the room. Although vestibular and somatosensory information could have been used to adjust the rod, visual context dominated orientation perception. This phenomenon is called the tilted-room illusion, and it dramatically illustrates the powerful contribution of visual information to orientation perception. The tilted-room illusion has been exploited in several roadside attractions and amusement parks (Banta, 1995; Shimamura & Prinzmetal, 2000). In these settings, people's misperception of vertical and horizontal is such that balls appear to roll uphill and people appear to stand at gravity-defying angles. Such compelling illusions have resulted in these settings being called "antigravity houses" (also see www.illusionworks.com). For over a hundred years, visual scientists have believed that visual illusions provide unique insights into normally adaptive visual processes. For example, Helmholtz (1881/1962) commented, The study of what are called illusions of the senses is a very prominent part of the senses; for just those cases which are not in accordance with reality are particularly instructive for the discovering the laws of those means and processes by which normal perception originates, (p. 251) With rare exception (e.g., Ward & Coren, 1976), most theories have attributed visual illusions to "information processing mechanisms that are normally adaptive" (Gregory, 1968, p. 66). The controversy arises as to what mechanisms are being reflected by particular illusions. For example, the Zollner illusion has been attributed to a variety of mechanisms, including spatial frequency coding in early vision (Ginsburg, 1984), inhibition among orientation tuned cells (Blakemore, Carpenter, & Georgeson, 1970), and the mechanisms responsible for linear perspective (e.g., Gillam, 1980; Green & Holye, 1963; Gregory, 1963). We offer a new explanation that also involves a normally adaptive process. We propose that the perception of line orientation (e.g., Zollner illusion), collinearity (e.g., Poggendorff illusion), line length (e.g., Ponzo illusion), and line curvature (e.g., Wiindt-Hering illusion) are caused by the same mechanisms that are responsible for the perception of vertical and horizontal. Because these mechanisms are related to people's ability to perceive a constant orientation despite changes in retinal orientation or even The rod-and-frame effect illustrates that a very sparse visual stimulus can also create orientation distortions (Witkin & Asch, 1948b). In the rod-and-frame task, observers adjust a rod to vertical or horizontal. The rod is surrounded by a tilted frame. Observers tend to adjust the rod in the direction of the tilted frame. Note that the rod-and-frame effect is local in that it is greater when the rod is close to the frame and is reduced in magnitude as the gap between the rod and frame increases (e.g., Zoccolotti, Antonucci, & Spinelli, 1993). The rod-and-frame effect and tilted-room illusion are similar to the tilt-induction effect (Gibson, 1937). Gibson presented stimuli similar to Figure 1A. Observers noted that a truly vertical line appeared to tilt in the direction opposite to the surrounding context lines (the apparent tilt is indicated, in exaggerated form, by the dashed line). Hence, when observers adjusted the central line to William Prinzmetal, Department of Psychology, University of California, Berkeley; Diane M. Beck, Department of Psychology, University College London, London, England. This research was supported in part by National Institutes of Health Grant MH51400. We thank Marty Banks, Art Shimamura, Steve Palmer, Edward Hubbard, and Nancy Kim for their help on this project. Correspondence concerning this article should be addressed to William Prinzmetal, Department of Psychology, University of California, Berkeley, California 94720-1650. Electronic mail may be sent to wprinz@socrates.berkeley.edu. 206 TILT CONSTANCY AND ILLUSIONS Figure 1. In all of the panels, the actual stimulus is shown in black, and the illusion, in exaggerated form, is shown with a dashed line. (A) The tilt-induction effect: Vertical lines are perceived to be tilted in the direction opposite the context lines. (B) The Zdllner illusion: The long oblique context lines in the tilt-induction figure have been replaced by many short context lines. (C) The standard Zollner illusion. (D) In the tilt-induction effect, Gibson (1937) observed that vertical context lines increased the apparent slope of an oblique line. (E) The tilt-induction effect from Panel D in a context more similar to the Poggendorff illusion. (F) The Poggendorff illusion can be considered a consequence of the tilt-induction effect in Panel D. vertical, they erred in the direction of the context lines, just as in the tilted-room illusion (also see Bouma & Andriessen, 1970; Kramer, 1978). When the context is presented before the stimulus, the effect is known as the tilt aftereffect (e.g., Campbell & Maffei, 1971; Gibson & Radner, 1937). The salient difference between the tilt-induction and rod-and-frame stimuli is the size of context: In the rod-and-frame effect, the stimuli typically subtend many tens of degrees of visual angle, whereas the tilt-induction experiment typically involves smaller stimuli. However, Coren and Hoy (1986) and Wenderoth and Johnstone (1988, Experiment 4) have obtained rod-and-frame effects with stimuli in the size range of the tilt-induction effect. We argue that the tilted-room illusion, the rod-and-frame effect, and the tilt-induction effect are caused by the same mechanisms (cf. Howard, 1982, pp. 155-156; Singer, Purcell, & Austin, 1970). Several investigators have speculated that the Zollner illusion is an instance of the tilt-induction effect (Day, 1972; Howard, 1982, 207 pp. 156-157). The similarity of the illusions can be seen in Figures 1A and IB, in which the flanking context lines in Figure 1A have been replaced by many shorter context lines in Figure IB. The Zollner illusion, as typically presented (Figure 1C), illustrates the effect of local context, in which the oblique lines on the left govern the slope of the left long line and the oblique lines on the right affect the slope of the right long line. Moreover, the tilt-induction effect and the Zollner illusion behave similarly with the manipulation of a host of independent variables (see Prinzmetal, Shimamura, & Mikolinski, in press). Many visual illusions, such as the Wundt-Hering illusion and Fraser spiral, can be understood as instances of the Zollner illusion (Fisher, 1968). It may be that the Cafe Wall illusion is also an instance of the Zollner illusion (Prinzmetal et al., in press). Historically and functionally, the Poggendorff illusion is also related to the Zollner illusion (Boring, 1942). In I860, ZSUner submitted his illusion for publication. The journal editor, Poggendorff, noticed that the oblique lines did not appear to be collinear. The functional relationship of the Poggendorff illusion to the tilt-induction effect is illustrated in Figures ID, IE, and IF. Gibson (1937) noted that, in the stimulus illustrated in Figure ID, the central oblique line appeared to be inclined at a larger angle than it actually was, illustrated again by the dashed line. (We have verified this in our laboratory.) If we simply displace the central line and its resulting illusory percept, as we did in Figure IE, we can see the similarities to the Poggendorff illusion in Figure IF. As a consequence of the misperception of the orientation of the central line (i.e., the tilt-induction effect), the oblique lines do not appear collinear. Note that it is not just that acute angles appear larger than they are, because the Poggendorff illusion can be created without any angles at all (e.g., see Day & Jolly, 1987; Pressey & Sweeney, 1972; Wilson & Pressey, 1976). We argue that the Poggendorff and Zollner illusions are caused by a misperception of orientation, not a misperception of angles. By our account, all of the geometric illusions described thus far can be related to orientation. Our approach can also account for other distortions, such as the line length observed in the Ponzo illusion. Figure 2 demonstrates how the Ponzo illusion can be considered a case of the Zollner illusion. In Figure 2A the vertical lines appear further apart at the top of the figure than at the bottom of the figure (Zollner illusion). Similarly, the horizontal line at the top of Figure 2B seems longer than the bottom horizontal line (Ponzo illusion). That is, the ends of the top horizontal lines appear Figure 2. The relation of the Ponzo and Zollner illusions. (A) The Z6Uner illusion with the Ponzo illusion suggested with bold lines. (B) The Ponzo illusion with the Zollner illusion in the background. 208 PRINZMETAL AND BECK further apart than the bottom horizontal lines. Figure 2A is drawn to suggest that the Ponzo illusion is part of the Zollner illusion, and in Figure 2B, the Ponzo is hidden in the Zollner illusion. We have conducted experiments on the tilt-induction effect with stimuli similar to those shown in Figures 3 A and 3B that compared the magnitude of the Ponzo illusion with the tilt-induction effect (Prinzmetal et al., in press). With stimuli such as those in Figure 3 A, observers adjusted the free end of the bottom horizontal line so that it appeared to be vertically aligned, according to gravity, with the end of the top horizontal line. The oblique line on the right caused a tilt-induction effect so that observers erred by making the line too long. The Ponzo illusion was measured with stimuli such as those in Figure 3B. Observers adjusted the length of the bottom horizontal line so that it matched the length of the top line. Because the Ponzo illusion involves a tilt-induction effect at either end, one would predict that the Ponzo illusion would be twice as great at the tilt-induction task, and this is what was found. Additionally, across observers, the correlation between the two illusions was .70. Prinzmetal et al. also compared predictions of several theories of the Ponzo illusion with the tilt-constancy theory. The results were consistent with the tilt-constancy theory. In summary, the tilt-constancy theory proposes that the tiltinduction effect and the Zollner, Poggendorff, and Ponzo illusions are due to a misperception of orientation caused by local visual context. Further, we propose that the mechanism responsible for the local misperception of location is the same as the mechanism that causes the tilted-room illusion and the rod-and-frame effect. The possibility that the Zollner illusion and the tilt-induction effects are caused by the same mechanism as the tilted-room illusion and the rod-and-frame effects has been the subject of controversy. On the one hand, Day (1972) suggested that the Zollner illusion is caused by the same mechanism as the tiltedroom illusion.1 On the other hand, Howard (1982, pp. 155-156) asserted that the "frame effects" (i.e., tilted room illusion, rod-andframe effect) are caused by a different mechanism than the "tiltnormalization" effects (i.e., Zollner illusion, tilt-induction effect). Heretofore, the evidence for either position is weak. Howard (1982) gave three reasons for believing that the frame and tilt-normalization effects are caused by different mechanisms. First, he asserted that the frame effects are many times larger than the normalization effects and therefore must be caused by different Figure 3. The Ponzo illusion considered as a consequence of the tiltinduction effect. In Panel A, the ends of the horizontal lines do not seem to be horizontally aligned. This effect makes the top horizontal line in Panel B appear longer than the bottom horizontal line. mechanisms. Strangely, he cited a misperception of orientation of 20°. We know of only one study that found errors of that magnitude. In very special circumstances, Asch and Witkin (1948) reported a tilted-room illusion of 20°. However, in their experiment, observers were seated in a chair that was tilted 24°. Furthermore, there was an elaborate procedure, lasting 7 min, to attempt to get observers to perceive the tilted room as upright. The magnitude of the tilted-room illusion is usually much less, and the rod-and-frame effect is typically even smaller (e.g., Wenderoth, 1974; Witkin & Asch, 1948b). In fact, Shimamura and Prinzmetal (2000) found that the tilted-room illusion can be of a similar magnitude as the tilt-induction effect. In general, the larger and more elaborate the context, the greater the illusion. This observation seems hardly surprising. Second, Howard (1982) claimed that tilted-frame effects are much more in evidence when the frames are in the periphery, but the tilt-induction effect is not greater in the periphery (Muir & Over, 1970). There are two pieces of evidence for this claim. First, a series of studies by Ebenholtz and his colleagues have shown that a larger frame leads to a larger illusion in the rod-and-frame task (e.g., Ebenholtz, 1977,1985; Ebenholtz & Callan, 1980). It is clear that as the frame becomes larger, it becomes more peripheral. However, these studies confound frame eccentricity and frame size. As we have commented, it does not seem surprising that a larger context would lead to a larger illusion.2 The second piece of evidence for the claim that the periphery is special is a study by DiLorenzo and Rock (1982). They conducted a rod-and-frame task with two frames, an inner frame and an outer (peripheral) frame. They found an effect of the outer frame but not the inner frame. However, the outer frame was always presented first, alone, for a period of 4 min, before the inner frame was presented. Hence, the procedure may have biased observers to use the outer frame. Finally, Howard (1982) claimed that the tilted-room illusion is affected by cognitive variables, whereas the tilt-induction effect (and other small-scale illusions) is not. By cognitive variables, Howard meant knowledge of the canonical orientation of objects such as flower vases and tables. However, there is no evidence that the tilt-induction effect is not influenced by cognitive variables. Furthermore, this argument is logically inconsistent in that Howard claimed that the tilted-room and rod-and-frame effects are caused by the same mechanism because they both involve large peripheral stimulation, whereas the tilt-induction effect does not. However, the rod-and-frame effect does not involve familiar objects (as in the tilted-room effect). Hence, by Howard's arguments, it is unclear whether these illusions should be classified on the basis of their scale, on whether they involve peripheral stimulation, or on whether they are influenced by cognitive variables. 1 We know of no one who has suggested that the Poggendorff and Ponzo illusions were caused by the misperception of orientation. Note that the misperception of orientation is not the same as a misperception of angles, a theory that is discussed in the General Discussion section. 2 Zoccolotti et al. (1993) replicated the finding that a larger frame leads to a larger rod-and-frame effect. As mentioned earlier, they found that with a given frame size, the size of the rod affected performance. As the rod increased in size (so that the gap between the rod and frame got smaller), the magnitude of the illusion increased. TILT CONSTANCY AND ILLUSIONS Day's (1972) position, that the tilted-room illusion is caused by the same mechanism as the Zollner illusion and the tilt-induction effect, is consistent with the tilt-constancy theory. The position is parsimonious in that a single mechanism is proposed for both phenomena. Furthermore, it provides a functional account of the tilt-induction illusion because the tilted-room illusion is related to the mechanism of tilt constancy. Nevertheless, there is no compelling evidence to support this view. If the tilted-room illusion and the tilt-induction effect are caused by the same mechanisms, then one would predict that an independent variable that critically affects the tilted-room illusion should also affect the tilt-induction effect. In the following experiments, we manipulated such a variable. In summary, the tilt-constancy theory claims that the same mechanism that causes the incorrect perception of orientation in the tilted-room illusion (e.g., Asch & Witkin, 1948) is responsible for a variety of illusions, including the Zollner, the Poggendorff, and Ponzo illusions as well as the tilt-induction effect. A variable that has been known to affect the tilted-room illusion is the physical orientation of the observer relative to gravity. Asch and Witkin (1948) found that seating observers in a chair that was tilted 24° significantly increased the magnitude of the tilted-room illusion (Asch & Witkin, 1948). Tilting the observer also increases the rod-and-frame effect (DiLorenzo & Rock, 1982; Witkin & Asch, 1948b). That is, observers tend to disregard gravity-related cues to orientation in favor of visual cues when tilted. The finding that observer orientation affects performance in the tilted room leads to a straightforward prediction: If the tilt-induction effect and the Zollner, Poggendorff, and Ponzo illusions are caused by the same mechanism as the tilted-room illusion, they should also increase when observers are seated in a tilted position. In the present experiments, we compared the magnitude of these illusions when observers were seated upright and when they were seated in a chair, tilted 30° about the observer's line of sight. A few previous investigators have measured various illusions with observers in postures other than upright with inconsistent and conflicting results (Campbell & Maffei, 1971; Day & Wade, 1969; Wallace & Moulden, 1973; Wolfe & Held, 1982). We believe that there were two critical factors when testing an effect of body orientation that were not controlled in previous research but were controlled in the present experiments. First, the visual environment must be adequately manipulated. In measuring the tilt-induction effect, if the frame on the computer monitor, or the surrounding room, is visible, then most of the visual environment is aligned with gravity even if the inducing lines are not. In our experiments, the observers were tested in the dark, and the stimuli consisted of luminous lines on a black background. To ensure that observers did not see the frame of the monitor, we required them to wear goggles with dark neutral density lenses. Second, because the perception of orientation persists over time (i.e., the tilt aftereffect), observers sat in the dark for 30 s before beginning each experiment. We do not believe that the tilt-constancy theory can explain all illusions, only those that are due to the misperception of orientation induced by visual context. Hence, we do not expect that tilting the observer will increase nonorientation illusions. The MiillerLyer illusion is a robust illusion, but we believe that it has little to do with the perception of orientation for three reasons. First, observers who evinced strong Zollner and Poggendorff illusions 209 were not those who showed a strong Miiller-Lyer effect (Coren, Girgus, & Erlichman, 1976). Second, as we mentioned earlier, the strength of many of these illusions varies with its orientation, and in our lab we have failed to find such an effect with the MullerLyer illusion. Finally, with the Zollner, Poggendorff, and Ponzo illusion, the orientation of the context lines critically determines the magnitude of the illusion. In contrast, with the Miiller-Lyer illusion, it is the distance the "wings" extend beyond the horizontal line that determines the illusion magnitude (Coren, 1986). Hence, we predicted that the Muller-Lyer illusion would not vary when observers were tilted. To ensure that a null finding was not the result of a floor or ceiling effect, we manipulated the strength of the Miiller-Lyer illusion, as described below. In Experiment 1, we tested the magnitude of the Zollner, Poggendorff, Ponzo, and Miiller-Lyer illusions, as well as the tilt-induction effect, with observers seated upright and tilted 30°. Asch and Witkin (1948) tilted observers 24° and Witkin and Asch (1948b) used 28°. We wanted to make sure that our manipulation was at least as strong as theirs, so we tilted our observers 30°. Experiment 2 was a control condition to test for the effect of tilting observers without visual context. Experiment 3 tested observers on the Zollner, Ponzo, and Poggendorff illusions when seated upright and tilted, while controlling for retinal orientation. Finally, in Experiment 4, we tested the effect of tilting observers on a second nonorientation illusion, specifically, an illusion of size induced by linear perspective. Experiment 1 Method Procedure. Each observer was tested in a single session that lasted about 1 hr. Half of the observers performed the tasks first seated upright, and then seated in a chair lilted 30° counterclockwise (from the observers' perspective), whereas the remaining observers were first tested seated in the tilted position. The order of the tasks within the first or second half of the experiment was counterbalanced across observers. The results from half of the observers on the Zollner task were lost because of a computeroperator error. Nevertheless, the order of conditions for the observers whose data were included was completely counterbalanced, as described above. Observers were tested for 24 trials in each task. In measuring the tilt-induction effect, observers moved the top dot back and forth so that it was vertically aligned, according to gravity, with the bottom dot (see Figure 4). Specifically, observers were told that if the two dots were tennis balls, the top dot should be placed so that if dropped, it would land squarely on the bottom dot in the real world. Observers used a joystick to move the top dot horizontally back and forth until they were satisfied with their setting, and then pressed a button on the joy stick to begin the next trial. Observers were told to take as much time as they wanted in making their settings. In the Zollner task (Figure 5A), observers moved the dot up and down, using the joystick, until it appeared to be collinear with the long line. The Poggendorff illusion (Figure 5B) was assessed by having observers move the bottom oblique line back and forth, horizontally, until it appeared collinear with the top oblique line. To measure the Ponzo illusion (Figure 5C), observers adjusted the length of the vertical line on the left until it appeared to be the same length as the line on the right. We used the Brentano version of the MUller—Lyer illusion. The observers' task was to adjust the location of the central arrowhead so that its tip would bisect the horizontal "shaft." There were two versions of the Miiller-Lyer illusion, as shown in Figure 5D. On 12 trials, the "wings" 210 PRINZMETAL AND BECK 1 degree Figure 4. The stimuli used to measure thetilt-inductioneffect. Observers moved the top dot, in the direction indicated by the arrow, to be vertically aligned with the bottom dot. The stimulus is drawn to scale, but in the actual experiment, the stimuli consisted of luminous lines on a black background. were long, and on 12 trials the wings were short. These were presented in a random order. It has been found that the length of the wings affects the strength of the illusion: The longer the wings, the greater the illusion (see Coren & Girgus, 1978, p. 31). We included this manipulation to ensure that a null result of tilting the observer was not due to the MUller-Lyer illusion being at floor or ceiling. Each task began with the observer sitting in the dark for 30 s, whether tilted or upright. The starting position of the adjustable component of the stimulus (e.g., vertical location of dot in the Zollner illusion) was randomly set before each trial. In addition, in the tilt-induction task, not only was the horizontal location of the top dot randomly set at the beginning of each trial, but the vertical position of the pair of dots was also randomly set within approximately ± 0.5° of visual angle. The reason for this random displacement was so observers would not use the relationship between the dot and a particular line in the grating to determine their setting. Stimuli. Observers were tested in a dark room while wearing goggles with dark neutral density filters. Hence, the stimuli appeared as luminous lines in an otherwise dark field. The stimuli were presented on a Macintosh computer, and the observers were seated at a viewing distance of 107 cm. The stimuli are illustrated in Figures 4 and 5. These figures are drawn to scale. The context lines in the tilt-induction effect stimulus (Figure 4) form an angle of 18° from vertical. Similarly, the oblique lines in the Zollner, Poggendorff, and Ponzo illusions were 18° from horizontal. In the MUllerLyer figure, the arrowheads formed an angle of 45°. In the tilt-induction effect (Figure 4), the distance between the centers of the dots subtended 2.84° of visual angle (160 screen pixels). The horizontal lines in the Zollner and Poggendorff illusions (Figures 5A and 5B) subtended 5.68° of visual angle (300 pixels). The oblique lines for the Zollner and Poggendorff illusions subtended 2.84° and 1.89° of visual angle, respectively (150 and 100 pixels). The length of the standard line in the Ponzo illusion subtended 2.84° of visual angle (150 pixels). Finally, the length of the horizontal line in the Milller-Lyer figure subtended 4.73° of visual angle (250 pixels), whereas the arrowheads subtended either 0.379° or 0.758° of visual angle (20 or 40 pixels). Apparatus. Observers were seated in a chair that could be set at upright or tilted 30° counterclockwise. The chair was created from a standard office furniture chair with armrests. The armrests of the chair were padded. The chair was fitted with a footrest and a shelf on which observers rested the joystick. Both the footrest and shelf tilted with the chair. Observers' heads wererestrainedby a chin rest and lateral support bars so that thenheads were at the same angle as the chair. Figure 5. The stimuli used to measure the (A) Zollner illusion, (B) Poggendorff illusion, (C) Ponzo illusion, and (D) Muller-Lyer illusion. The arrows indicate the adjustments that observers could make. The stimuli are drawn to scale, but in the actual experiment, the stimuli consisted of luminous lines on a black background. TILT CONSTANCY AND ILLUSIONS Observers. Twenty observers participated in the experiment, half of them were male and half female. However, because of the abovementioned computer error, the results from only 10 observers were included in the Zollner task. The average age of the observers who participated in the experiment was 22 years. All reported that they had normal or corrected-to-normal vision with no known visual deficits. Observers were recruited from among the undergraduate population of the University of California, Berkeley. Results The tilt-induction effect and the Zollner, Poggendorff, and Ponzo illusions were all dramatically and significantly increased when the observers were tilted 30° compared with when they were in the upright condition (see Table 1). The dependent variable for the tilt-induction effect was the deviation from vertical of a virtual line connecting the top and bottom dots, in degrees, with positive deviations in the direction of the grating (see Figure 4). For the Zollner and Poggendorff illusions (see Figure 5), the dependent variable was the difference, in screen pixels, from the average adjusted height of the dot to the correct (collinear) location. (Note that 52.8 pixels are 1° of visual angle.) The magnitude of the Ponzo illusion was measured as a percentage: the length of adjustable line divided by the length of the standard line. As shown in Table 1, tilting the observer significantly increased each of these illusions. Table 1 also indicates the average standard deviation for each illusion, that is, the standard deviation over the 24 trials for each condition, averaged over observers. An additional analysis was conducted with the Poggendorff illusion. Several investigators (e.g., Weintraub & Krantz, 1971; Weintraub, Krantz, & Olson, 1980) have suggested that the version of the Poggendorff illusion that we used should be measured in degrees. That is, the differences, in degrees, between the angle of the oblique line (i.e., 18° in this experiment) and the angle formed by a virtual line connecting the dot and the intersection of the oblique line and horizontal line. This dependent variable has the advantage that it is invariant to viewing distance. Therefore, we performed an additional analysis on the Poggendorff illusion in which we first transformed each response into degrees error. The illusion was significantly greater for the tilted than upright conditions, 8.45° versus 6.01°, F(l, 19) = 9.66, p < .05. In contrast, the Miiller-Lyer illusion was not affected by the observer orientation (see Table 1). The dependent variable for the Muller-Lyer illusion was the average distance, in screen pixels, Table 1 Results From Experiment 1 Effect Upright condition Tilted condition df F 71.72** 96.63** Tilt-induction effect Zollner illusion Poggendorff illusion Ponzo illusion 2.4° (3.0) 7.3 pixels (4.5) 9.4° (4.9) 26.4 pixels (7.3) 1,19 1,9 28.7 pixels (9.0) 5.9% (3.9) 38.8 pixels (11.3) 8.2% (4.4) 1,19 20.86** 1, 19 7.28* Muller-Lyer illusion 18.2 pixels (3.4) 19.0 pixels (3.7) 1,19 Note. Average standard deviations are in parentheses. *p < .05. **p < .01. 0.95 211 from the true bisection point. (Note that the "shaft" was 250 pixels in length.) There was no significant effect of observer orientation, but the magnitude of the illusion was significantly greater with long fins than short fins, 22.16 versus 14.98 pixels, F(l, 19) = 103.84, p < .01. The interaction of fin length and observer orientation was not significant, F(l, 19) = .05. For the upright condition, the average illusion for long and short fins was 21.72 and 14.64 pixels, respectively. For the tilted condition, the average illusion for long and short fins was 22.63 and 15.31 pixels, respectively. In summary, each illusion for which the tilt-constancy theory has an account (tilt induction, Zollner, Poggendorff, and Ponzo) behaved like Asch and Witkin's (1948) tilted-room illusion: The illusion increased when observers were tilted. However, tilting observers does not increase every illusion: It had no effect on the Miiller-Lyer illusion. We conducted a partial replication of this experiment with slightly different stimuli (e.g., the Zollner figure had two horizontal lines) and 10 additional observers with identical results: a significant effect of observer tilt with the Zollner, Ponzo, and Poggendorff illusions but no influence on the MiillerLyer illusion. Finally, we ran an analysis of the Miiller-Lyer illusion that combined the main experiment and the replication, so that there were 30 observers, and the effect of body tilt was still not reliable. Experiments 2 and 3 address two alternative accounts of this effect. Experiment 2 The dramatic effect of tilting the observer on the tilt-induction effect could be the result of tilting the observer per se and not the result of visual context on perception. It is known that tilting an observer 45° or more in the dark causes a line to appear tilted in the opposite direction. This effect is called the Aubert effect or A effect (Howard, 1982, p. 427; Witkin & Asch, 1948a), and it could account for our findings with the tilt-induction effect. However, tilting the observer less than about 30° causes a line to appear tilted in the same direction as the observer (E effect) and would run counter to our results. To test the effect of tilting the observer on orientation without context, we repeated the tilt-induction task of Experiment 1 but without the tilted grating (see Figure 1). The stimulus consisted of just two dots, and 20 observers were run in the tilted condition only. The average setting was 0.87° in the direction opposite the observer's body tilt. This value was not significantly different from 0, f(19) = 0.98, ns, and is in the opposite direction of that found in Experiment 1, which included a visual context (i.e., grating). The average standard deviation was 2.14°. Thus, the increase of the tilt-induction effect with tilting observers in Experiment 1 was not the result of the A effect but rather was due to an increased effect of visual context. Experiment 3 It has been known since the early days in the study of visual illusions that the Zollner and Poggendorff illusions are affected by the orientation of the stimulus (e.g., Judd & Courten, 1905; Velinsky, 1925). For example, Judd and Courten found that the magnitude of the Zollner illusion was less in the orientation 212 PRINZMETAL AND BECK illustrated in Figure 5 than when rotated 45°. Hence, the effect of tilting the observer in Experiment 1 might simply be an effect of rotating the stimulus on the retina rather than rotating the observer per se. To test for this possibility, we replicated Experiment 1 with the Zollner, Ponzo, Poggendorff, and Muller-Lyer tasks with the stimulus at the same retinal orientation when the observer was seated upright and when the observer was seated tilted 30°. We did not run the tilt-induction effect because we were not sure what to expect if the orientation of the grating was changed withrespectto retinal orientation because the observer's task was to indicate gravitational vertical. Table 2 Results From Experiment 3 Method orientation, tilting the observer still increased the magnitude of these illusions. We had previously run a pilot experiment with just the Zollner illusion with 10 observers, correcting forretinalorientation, and obtained the same results: The illusion was greater when observers were tilted even when retinal orientation was controlled. In this pilot experiment, the viewing distance was the same as in Experiments 1 and 2, and the Zollner stimulus was rotated 45° (like the Poggendorff stimulus in the present experiment). Note that the magnitude of the effect of tilting the observer was less than in Experiment 1 (compare Tables 1 and 2). Hence, it may be that there is some effect of retinal orientation. Alternatively, the difference between the magnitude of the effects in Experiments 1 and 3 may be due to the perceived orientation of the figures, not the retinal orientation. In an ingenious study, Spivey-Knowlton and Bridgeman (1993) varied the orientation of a Poggendorff stimulus and found that the magnitude of the illusion varied with orientation, producing a function similar to previous studies (e.g., Green & Hoyle, 1964; Greene & Pavlow, 1989; Leibowitz & Toffey, 1966). However, when they surrounded the stimulus with a luminous frame of different orientations, it was the orientation of the frame that was critical, not the retinal orientation. The results for the Miiller-Lyer illusion surprised us. The illusion was significantly less when observers were tilted than when they were upright (see Table 2). The magnitude of the effect was small (18.5 vs. 16.9 pixels) but reliable. As discussed above, we previously did not obtain any effect of tilting the observer on the Miiller-Lyer illusion. Hence, it is possible that this result is a Type I error. Nevertheless, note that tilting the observer had the opposite effect on the Muller-Lyer illusion as the other illusions. This result is consistent with claim that the Miiller-Lyer illusion is caused by a different mechanism than the other illusions that we tested. Procedure. Half of the observers began with the upright condition, and the other half began with the tilted condition. Observers were tested for 30 trials in each condition. For the upright condition, the stimulus was identical to Figure 5 except that the Poggendorff illusion was rotated 45° in a counterclockwise direction from that shown in Figure 4B. We used a 45° orientation, because depending on the observers' eye rotation (see below), further rotating the stimulus in the tilted condition made the short lines horizontal with respect to gravity for some observers but not for others. We wanted to keep the stimulus similar for all observers regardless of their eye rotation. Also, for the Miiller-Lyer illusion, we used only the short wing version (see Figure 5). When the observer was tilted 30°, the stimulus was further rotated so that it would form the same retinal image as when the observer was upright. When observers are tilted, their eyes rotate in their orbits in the direction opposite to their body rotation (Howard, 1982, p. 383), and the amount of eye rotation varies from observer to observer. Thus, we measured each observer's eye rotation (when seated tilted 30°) and adjusted the stimulus so that it had the same retinal orientation when the observer was upright and tilted. Rotational eye movements were measured in the following manner. At the beginning of the session, observers were seated in a normal chair in a standard upright posture. They looked at a photographic strobe light that was masked with a vertical slit. When the strobe flashed, it left an afterimage of a line that was retinally vertical. The observers then quickly moved to the tilted chair. Next, observers adjusted the orientation of a line on the computer monitor until it lined up perfectly with his or her afterimage. If there was no eye rotation, then observers would set the line at 30°, the angle of body tilt. If observers' eyes rotated back to vertical, then they would set the line to vertical (0°). Thus, eye rotation was simply the difference between the angle that observers adjusted the line and their body tilt. Rotational eye movements ranged from 0° to 13°, across observers, and averaged 4.5°. No observer reported difficulty in the eye rotation task. We checked the reliability of this method with 2 observers on several different occasions and measurements agreed within 1°. The viewing distance was 81 cm instead of the 107 cm used in Experiments 1 and 2? Hence, the horizontal lines in the Zollner and Poggendorff illusions subtended 7.4° of visual angle instead of the 5.68° in Experiment 1. Thus, all dimensions of the stimuli subtended approximately 75% of those in Experiment 1. Observers. Twenty-four observers, recruited from among the undergraduate population of the University of California, Berkeley, participated in the experiment. The average age was 19 years; 19 were female and 5 were male. Effect Upright condition Tilted condition F(\, 23) Zollner illusion Poggendorff illusion Ponzo illusion 12.2 pixels (6.9) 35.6 pixels (11.2) 6.9% (4.3) 20.2 pixels (8.6) 40.1 pixels (10.6) 8.5% (4.4) 24.73** 4.51* 5.77* Muller-Lyer illusion 18.5 pixels (4.0) 16.9 pixels (4.0) 5.19* Note. Average standard deviations are in parentheses. *p<.05. **p<.01. Experiment 4 In Experiment 1, the illusions that we claim are related to orientation increased when the observer was tilted (Zollner, Ponzo, Poggendorff, and tilt-induction illusions). We used the MiillerLyer illusion as an example of a nonorientation illusion, and its magnitude was unchanged by tilting the observer. Because the causes of the Miiller-Lyer illusion are uncertain, however, we Results and Discussion The magnitude of the Zollner, Poggendorff, and Ponzo illusions were significantly greater in the tilted condition than in the upright condition (see Table 2). Thus, even when we controlled for retinal 3 Experiment 3 was conducted about 2 years after the other experiments reported in the article. The chair apparatus and the arrangement of the laboratory furniture had changed during that period, resulting in a change in viewing distance. TILT CONSTANCY AND ILLUSIONS 213 Figure 6. The stimulus used in Experiment 4. The stimulus is drawn to scale, but in the actual experiment, the stimuli consisted of luminous lines on a black background. wanted to test a control condition that was clearly not caused by the misperception of orientation. The size-constancy illusion shown in Figure 6 provides an interesting contrast to what we claim are orientation-based illusions.4 Most observers perceive the cylinder in the middle of the figure to be taller than the cylinder at the bottom of the figure. This effect arises because linear perspective and other cues cause the cylinder in the middle of the figure to appear further away than the cylinder at the bottom. If two objects subtend the same visual angle, the object that appears further away will be perceived as larger. We do not think that this effect is related to orientation, so tilting the observer should not affect this illusion. However, several theories account for the Ponzo, Zollner, and Poggendorff illusions in terms of linear perspective or perceived depth (e.g., Gillam, 1980; Green & Holye, 1963; Gregory, 1963). Linear perspective does not change with the retinal orientation of the stimulus (Gregory, 1964), but we do not know how tilting the observer would affect linear perspective. Tilting the observer increases the Ponzo, Zollner, and Poggendorff illusions. If these are caused by the mechanisms responsible for linear perspective and size constancy, then tilting the observer might also increase the illusion illustrated in Figure 6. Method Procedure. Observers were presented with stimuli such as those illustrated in Figure 6. The task was to adjust the height of the cylinder at the bottom to match the height of the cylinder in the middle. Observers adjusted the height with a joystick and were under no time pressure. The adjustments that observers made only affected the height of the bottom cylinder and left its width unaffected (see Stimulus section). Each observer made 24 adjustments in the tilted condition and 24 adjustments in the upright condition. Half of the observers were tested upright first, and half were tilted first. Twenty observers, selected as before, participated in a single session that lasted about 10 min. Stimulus. The stimulus is shown in Figure 6. The figure is drawn according to the rules of linear perspective (see Gillam, 1981). It appeared as luminous and gray areas (the guitars) on a black background. Because a line drawing using linear perspective can be ambiguous, the guitars were added to disambiguate the depth relations. For example, without the guitars and sign, Figure 6 could represent a pyramid viewed from above. The sign (Home, Sweet Home) was added for artistic balance. As in the previous experiments, the stimuli were presented on a Macintosh computer, and the observers were seated at a viewing distance of 107 cm. At this distance, the area covered by the stimulus subtended 15.2° X 9.5° of visual angle (802 X 500 screen pixels). The standard cylinder (in the center) subtended 8.3° in height (44 pixels) and 6.1° in width (32 pixels). The initial height of the adjustable cylinder (on the bottom) was randomly set from 20 to 60 pixels, but the width did not change. The change in height was accomplished by stretching or contracting the rectangle on which the cylinder was drawn in a vertical direction only. (Note that the width of the cylinder might be affected by a Ponzo-like illusion caused by the oblique lines. For this reason, the manipulation was only in the vertical direction.) In all other respects, this experiment was similar to Experiment 1. Results and Discussion Every observer but 1 adjusted the cylinder too tall (n = 20). The error can be expressed as a percentage of the height of the standard (as with the Ponzo illusion). Observers adjusted the cylinder 14.0% too high in the upright condition and 12.3% too high in the tilted condition. Note that this difference, though not reliable, F(l, 19) = 1.21, is in the opposite direction as the effect of tilting the observer on the orientation illusions in Experiment 1. The standard deviations, averaged over the 20 observers, were 5.1% and 6.3% for upright and tilted conditions, respectively. 4 We are grateful to Jeremy Wolfe for suggesting this experiment. PRINZMETAL AND BECK 214 These results would seem to be fairly damaging for accounts of the Ponzo, Poggendorff, Zollner, and tilt-induction illusions that make reference to linear perspective. Tilting the observer increases the tilted-room illusion (Asch & Witkin, 1948), the rod-and-frame effect (DiLorenzo & Rock, 1982; Witkin & Asch, 1948b), as well as the illusions tested in Experiment 1. However, tilting had no effect on linear perspective and size constancy. Of course it is difficult to prove the null hypothesis. One might worry, for example, that the effect of perspective was too small, leading to a floor effect. Note that the effect of perspective on the cylinder height was similar in magnitude as the Ponzo illusion in Experiments 1 and 2 (see Tables 1 and 2). Both are illusions of linear extent, in a vertical direction. Importantly, there is no a priori reason to believe that tilting the observer, or the stimulus, would affect linear perspective. Gregory (1964), for example, asserted that "when a perspective figure is rotated, the perspective does not change" (p. 303). Nevertheless, it was important to demonstrate this dissociation between the effect of tilting the observer in the orientation illusions and an illusion based on linear perspective. There is one account that may allow linear perspective (and other theories) to play a role in some of these illusions, however. It may be that some of these illusions have multiple causes, as suggested by Coren and Girgus (1978). For example, it could be that the Poggendorff illusion is caused by both tilt constancy and linear perspective. The present experiments only manipulated the tilt-constancy component, but linear perspective (or other mechanisms) may still play a role. The present experiments were designed to test the tilt-constancy explanation and to provide an empirical link between the largescale illusions (tilted-room and rod-and-frame effects) and the orientation illusions. To rule out other theories, one needs a situation in which competing theories make different predictions. We have tested various theories in this manner with the Ponzo illusion (Prinzmetal et al., in press). With the Ponzo illusion, Prinzmetal et al. demonstrated that the tilt-constancy theory accounted for 100% of the Ponzo illusion, leaving little role for other mechanisms. Furthermore, Prinzmetal et al. demonstrated that the following theories were neither necessary nor sufficient to account for the Ponzo illusion: (a) the low-pass filter theory (Ginsburg, 1984), (b) the assimilation theory (Pressey & Epp, 1992), (c) a sizecomparison theory (Kunnapas, 1955), (d) the pool-and-store theory (Girgus & Coren, 1982), and (e) the linear perspective and size-constancy theory (e.g., Gillam, 1980; Gregory, 1963). More research will be necessary to completely rule out the possibility that these theories play a role for some of the other illusions. General Discussion We hypothesized that the Zollner, Poggendorff, and Ponzo illusions and the tilt-induction effect were caused by the same mechanism that causes the tilted-room illusion. Tilting an observer increases the tilted-room and the rod-and-frame effects (Asch & Witkin, 1948; DiLorenzo & Rock, 1982; Witkin & Asch, 1948b). If the same mechanisms are responsible for these illusions, then tilting the observer should also increase these illusions, and it did. The discussion that follows is organized around three issues: First, we discuss our findings in relation to our theory versus previous theories of these illusions. Second, we suggest an algorithm for tilt constancy and discuss its neural implementation. Finally, we briefly consider a few empirical issues left unresolved by the present research. Our present results are inexplicable by previous theories of these illusions, which include those based on linear perspective (e.g., Gillam, 1980; Green & Holye, 1963; Gregory, 1963), the low-pass filter theory (Ginsburg, 1984), and inhibition among orientationtuned cells (Blakemore et al., 1970). Unlike the tilt-constancy theory, none of these theories predict an increase in the magnitude of these illusions when an observer is tilted. The present research suggests that the relevant mechanisms revealed by these illusions are those that determine orientation, and in particular, local visual cues to orientation. Of course, as we pointed out previously, we cannot rule out the possibility that other mechanisms may play some role in these illusions, but any theory of these illusions must contain a reference to the perception of orientation. Our account of these results, the tilt-constancy theory, has some similarities with theories that appeal to size constancy and linear perspective (e.g., Day, 1972; Gillam, 1980; Gregory, 1963). There are several distinct variants of this theory (see Green & Hoyle, 1964), but all appeal to mechanisms related to size constancy. Size constancy is, of course, the ability to perceive objects in their true size regardless of distance. Tilt constancy is the ability to perceive objects in their true orientation regardless of the orientation of the observer. Hence, both theories appeal to normally adaptive processes. Furthermore, although there are a number of determinants of perceived distance, the theories that account for the visual illusions treated in this article only appeal to linear perspective. Similarly, there are several determinants of perceived orientation in addition to visual information, such as vestibular and somatosensory information, but we only appeal to the misperception of orientation due to visual information. Note that both the size-constancy family of theories and the tilt-constancy theory account for a wide range of illusory phenomena but not precisely the same phenomena. For example, we have no account of the Miiller-Lyer illusion, whereas the Miiller-Lyer illusion is often used as the foremost example for the sizeconstancy theories. Indeed, it is not a virtue for the same theory to account for both the Miiller-Lyer illusion and the other illusions because, as we pointed out, both previous research and our present findings suggest that they are not caused by the same mechanisms. To this previous evidence, we add that the Zollner, Poggendorff, and Ponzo illusions and tilt-induction effect behaved differently than the Miiller-Lyer illusion when observers were tilted. Hence, although size constancy may provide a good explanation of the Miiller-Lyer illusion (see e.g., Gregory & Harris, 1975), it does not account for our results, and Experiment 4 clearly demonstrates that size constancy behaves differently than the orientation-based illusions. Our theory should not be confused with another theory that acute angles are overestimated (Hering, 1861). This theory has been applied to the Zollner and Poggendorff illusions and the tilt-induction effect. It has not been applied to the Ponzo illusion. In modern physiological terms, the overestimation of angles is claimed to be a consequence of the inhibition of neurons in VI (e.g., Blakemore et al., 1970). According to this explanation, when two lines forming an acute angle are presented to an observer, each line activates a population of cells. The cells activated are those tuned to the orientation of each line. However, cells tuned to TILT CONSTANCY AND ILLUSIONS orientations similar to the lines are also activated. Cells activated by both lines (i.e., those with orientations between the lines) mutually inhibit each other. Thus, the total activation is shifted so that the angle is perceived as larger than it is. There is now ample evidence that the acute-angle theory is inadequate to account for the Zollner and Poggendorff illusions because these illusions can be obtained without any "angles" in the stimulus (e.g., Pressey & Sweeney, 1972; Wilson & Pressey, 1976). Indeed, our version of the tilt-induction effect in Experiments 1 and 2 (see Figure 4) does not contain lines that would mutually inhibit each other. Indeed, Tyler and Nakayama (1984) provided an example of a Zollner illusion with only lines of one orientation, and Day and his colleagues (Day & Jolly, 1987; Day, Jolly, & Duffy, 1987; Day, Watson, & Jolly, 1986) described a version of the Poggendorff illusion, using a bisection task and only two parallel lines (or even three dots). Furthermore, a theory based on simple inhibition between line segments cannot explain the results of tilting the observer. We are not suggesting that a theory based on neural architecture is inappropriate; later, we outline the beginning of a neural theory of perceived orientation. However, a theory based on the misperception of angles (as opposed to orientation) does not account for existing data. A misperception of orientation may cause a misperception of angles, but the cause of the illusions, according to the tilt-constancy theory, has its roots in the perception of orientation. A final family of theories, referred to as assimilation theories, has been used to account for many of these same illusions. These theories are diverse and include the low-pass filter theory of Ginsburg (1984), the assimilation theory of Pressey and his colleagues (Pressey, 1971; Pressey, Butchard, & Scrivner, 1971; Pressey & Epp, 1992), and the pool-and-store model of Girgus and Coren (1982). These theories also have no account of our present findings. Indeed, no theory that only appeals to visual mechanisms—as opposed to a mechanism of spatial representation—can account for the effects of tilting the observer. A complete theory of these illusions should specify the neural mechanisms underlying tilt constancy. Our results suggest that the mechanisms are likely to involve neurons that are tuned to orientation relative to the visual environment, rather than the observer's retinal orientation. In animal studies, Sauvan and Peterhaus (1995) searched in the cortex of awake macaque monkeys for cells whose tuning functions were related to the environment as opposed to the animal's orientation. They found virtually no cells in the striate cortex that showed this property. All of the cells in the striate cortex were tuned in relation to retinal coordinates. However, 40% of cells in extrastriate areas (Areas V2, V3, and V3a) were tuned in relation to environmental, not retinal coordinates. Likely candidates for the neural substrate of these illusions will be areas with an abundance of cells that are tuned to environmental coordinates. There are probably many ways of implementing tilt constancy with simple neural hardware. Environmentally tuned cells could be created from retinally tuned cells with a simple two-layer system. Suppose there is a population of retinally tuned vertical detectors in the first level. These cells will fire to vertical lines, but also to lines near vertical. If the stimulus contains lines that are near vertical, a second level of the system would be erroneously signaled that a vertical was present. The same process would happen with stimulus lines near horizontal. Thus, contours in the visual environment near retinal vertical would define orientation. Note 215 that this mechanism is distinct from lateral inhibition and low-pass filtering. However, like these theories, it invokes simple neural processes. A simple mechanism such as this might underlay what Gibson (1937) meant by a process of normalization. Of course, such a simple scheme is incomplete because it does not include gravity-based cues to orientation (i.e., vestibular and somatosensory cues). Our findings, along with previous findings by Asch and Witkin (1948) and others (e.g., DiLorenzo & Rock, 1982; Goodenough, Oltman, & Sigman, 1981; Templeton, 1973), suggest that when an observer is not in an upright position, visual cues play a greater role. There are many possibilities as to how visual and extravisual information are integrated. For example, it might be that the perceived orientation of a stimulus is a weighted linear average of retinal and gravity-based cues (Matin & Fox, 1989). Alternatively, the integration of information may be nonlinear (e.g., Carlson-Radvansky & Logan, 1997). Furthermore, there are many possible sites in the brain for the integration of visual and other information. In addition to the general theoretical issue discussed above, tilt-constancy theory raises a plethora of research issues, of which we mention only three. First, we do not have a precise description of why tilting an observer increases the tilted-room illusion and related illusions. In the most general sense, tilting the observer increases one's reliance on visual as opposed to gravity-based information. However, this description raises a number of empirical issues. For example, tilting the observer may increase the orientation illusions because it creates a conflict among retinal, visual, and gravity-based cues to orientation. If this hypothesis is correct, only body positions that create a conflict will increase these illusions. However, perhaps any body position that is not aligned with visual vertical will influence performance. In this case, a supine posture would also increase the tilt-induction effect and the Zollner, Ponzo, and Poggendorff illusions. Because the rod-and-frame effect is greater when observers are in a supine posture (Goodenough et al., 1981; Templeton, 1973), we would predict that supine observation would similarly affect the other illusions. We are currently testing this hypothesis. A second issue relates to the perception of pitch. There is a well-known pitch-box illusion that is analogous to the tilted-room illusion. Observers' perceived eye level is affected by the pitch of the visual environment. If observers are looking down a hall that is pitched down, for example, they will indicate that eye level is lower than it actually is (Kleinhans, 1970; Matin & Fox, 1989; Matin & Li, 1992,1996; Stoper, 1998; Stoper & Cohen, 1986). An issue that concerns us is whether the mechanisms of pitch perception (e.g., perception of eye level) are the same as the mechanisms for the perception of vertical and horizontal. Finally, the concept of reference frames plays a large role in other aspects of cognitive psychology, including recognition (Mermelstein, Banks, & Prinzmetal, 1979; Rock, 1974; Yin, 1969), object description, and attention (e.g., Behrmann & Tipper, 1999; Robertson, 1995). An important issue is whether the local frames of reference that cause distortions such as the Ponzo and Zollner illusions are the same as these other reference frames. To the extent that they are the same, manipulations that affect the illusions (e.g., tilting the observer) should have similar effects in these other domains. We proposed that the Ponzo, Zollner, and Poggendorff illusions and the tilt-induction effect are due to the visual mechanisms that PRINZMETAL AND BECK 216 create illusions of orientation in the tilted-room illusion (tiltconstancy theory). In the tilted-room illusion, tilting the observer increases the effect of visual context in relation to gravitational information. In a similar manner, tilting the observer also increases the magnitude of these illusions. The tilt-constancy theory broadens the functional significance of these illusions. These illusions are not only relevant to visual processes but also informative about the nature of human spatial representations. References Asch, S. E., & Witkin, H. A. (1948). Studies in space orientation: 0. Perception of the upright with displaced visual fields and with body tilted. Journal of Experimental Psychology, 38, 455-477. Banta, C. (1995). Seeing is believing: Haunted shacks, mystery spots, and other delightful phenomena. Agoura Hills, CA: Funhouse Press. Behrmann, M., & Tipper, S. P. (1999). Attention accesses multiple reference frames: Evidence from visual neglect. Journal of Experimental Psychology: Human Perception and Performance, 25, 83-101. Blakemore, C , Carpenter, R. H., & Georgeson, M. A. (1970). Lateral inhibition between orientation detectors in the human visual system. Nature, 228, 37-39. Boring, E. G. (1942). Sensation and perception in the history of experimental psychology. New York: Appleton-Century-Croft. Bouma, A., & Andriessen, J. J. (1970). Inducted changes in the perceived orientation of line segments. Vision Research, 10, 333-349. Campbell, F. W., & Maffei, L. (1971). The tilt after-effect: A fresh look. Vision Research, 11, 833-840. Carlson-Radvansky, L. A., & Logan, G. D. (1997). The influence of reference frame selection on spatial template construction. Journal of Memory and Language, 37, 411-437. Coren, S. (1986). An efferent component in the visual perception of direction and extent. Psychological Review, 93, 391-410. Coren, S., & Girgus, J. S. (1978). Seeing is deceiving: The psychology of visual illusions. Hillsdale, NJ: Erlbaum. Coren, S., Girgus, J., & Erlichman, H. (1976). An empirical taxonomy of visual illusions. Perception <£ Psychophysics, 20, 129-137. Coren, S., & Hoy, V. S. (1986). An orientation illusion analog to the rod and frame: Relational effect in the magnitude of the distortion. Perception & Psychophysics, 39, 159-163. Day, R. H. (1972). Visual spatial illusions: A general explanation. Science, 175, 1335-1340. Day, R. H., & Jolly, W. J. (1987). A note on apparent displacement of lines and dots on oblique parallels. Perception & Psychophysics, 41, 187-189. Day, R. H., Jolly, W. J., & Duffy, F. M. (1987). No evidence for apparent extent between parallels as the basis of the Poggendorff effect. Perception & Psychophysics, 42, 561-568. Day, R. H., & Wade, N. J. (1969). Thereferencefor visual normalization. American Journal of Psychology, 82, 191-197. Day, R. H., Watson, W. L., & Jolly, W. J. (1986). The Poggendorff displacement effect with only three dots. Perception & Psychophysics, 39, 351-354. DiLorenzo, J., & Rock, I. (1982). The rod and frame effect as a function of the righting of the frame. Journal of Experimental Psychology: Human Perception and Performance, 8, 536—546. Ebenholtz, S. M. (1977). Determinants of the rod and frame effect: The role of retinal size. Perception & Psychophysics, 22, 531-538. Ebenholtz, S. M. (1985). Absence of relational determination in the rodand-frame effect. Perception & Psychophysics, 37, 303-306. Ebenholtz, S. M, & Callan, J. W. (1980). Modulation of the rod and frame effect: Retinal size vs. apparent size. Psychological Research, 42, 327334. Fisher, G. H. (1968). An experimental comparison of rectilinear and curvilinear illusions. British Journal of Psychology, 59, 23-28. Gibson, J. J. (1937). Adaptation, after-effect, and contrast in the perception of tilted lines: II. Simultaneous contrast and the areal restriction of the after-effect. Journal of Experimental Psychology, 20, 553-569. Gibson, J. J., & Radner, M. (1937). Adaptation, after-effect and contrast in the perception of tilted lines: I. Quantitative studies. Journal of Experimental Psychology, 20, 453-467. Gillam, B. (1980). Geometrical illusions. Scientific American, 242, 102111. Gillam, B. (1981). False perspective. Perception, 10, 313-318. Ginsburg, A. P. (1984). Visual form perception based on biological filtering. In L. Spillmann & B. R. Wotten (Eds.), Sensory experience, adaptation, and perception: Festschiriftfor Ivo Kohler (pp. 53-72). Hillsdale, NJ: Erlbaum. Girgus, J. S., & Coren, S. (1982). Assimilation and contrast illusions: Differences in plasticity. Perception & Psychophysics, 32, 555-561. Goodenough, D. G., Oltman, P. K., & Sigman, E. (1981). The rod-andframe illusion in erect and supine observers. Perception & Psychophysics, 29, 365-370. Green, R. T., & Holye, E. M. (1963). The Poggendorff illusion as a constancy phenomenon. Nature, 200, 611-612. Green, R. T, & Hoyle, E. M. (1964). The influence of spatial orientation on the Poggendorff illusion. Acta Psychologica, 22, 348-366. Greene, E., & Pavlow, G. (1989). Angular induction as a function of contact and target orientation. Perception, 18, 143-154. Gregory, R. L. (1963). Distortion of visual space as inappropriate constancy scaling. Nature, 199, 678-680. Gregory, R. L. (1964). Response to Brown and Houssiaadas. Nature, 204, 302-303. Gregory, R. L. (1968). Visual illusions. Scientific American, 219, 66-76. Gregory, R. L., & Harris, J. P. (1975). Illusion-destruction by appropriate scaling. Perception, 4, 203-220. Helmholtz, H. von. (1925). Handbuch der Physiologischen Optik [Treatise on physiological optics]. Lancaster, PA: Optical Society of America. Helmholtz, H. V. (1962). Popular lectures on scientific subjects (M. Kline, Ed., & E. Atkinson, Trans.). New York: Dover. (Original work published 1881) Hering, E. (1861). Beitrage zur Psychologie [Contributions to psychology]. Leipzig, Germany: Engleman. Howard, I. P. (1982). Human visual orientation. New York: Wiley. Howard, I. (1998, December). Knowing which way is up. Paper presented at the Vision Science Symposium, University of California, Berkeley. Howard, I. P., & Childerson, L. (1994). The contribution of motion, the visual frame, and visual polarity to sensations of body tilt. Perception, 23, 753-762. Judd, C. H., & Courten, H. C. (1905). The Zollner illusion. Psychological Review Monograph Supplements, 7 (Whole No. 27), 112-141. Kleinhans, J. L. (1970). Perception of spatial orientation in sloped, slanted, and tilted visual field. Unpublished doctoral dissertation, Rutgers University of New Jersey. Kramer, K. S. (1978). Inhibitory interactions as an explanation of visual orientation illusions. Unpublished doctoral dissertation, University of California, Los Angeles. Kunnapas, T. M. (1955). Influence of frame size on apparent length of a line. Journal of Experimental Psychology, 50, 168-170. Leibowitz, H., & Toffey, S. (1966). The effect of rotation and tilt on the magnitude of the Poggendorff illusion. Vision Research, 6, 101-103. Matin, L., & Fox, C. R. (1989). Visually perceived eye level and perceived elevation of objects: Linearly additive influences from visual field pitch and from gravity. Vision Research, 29, 315-324. Matin, L., & Li, W. (1992). Visually perceived eye level: Changes induced by a pitched-from-vertical 2-lin visual field. Journal of Experimental Psychology: Human Perception and Performance, 18, 257-289. Matin, L., & Li, W. (1996). Visually perceived eye level is influenced TILT CONSTANCY AND ILLUSIONS identically by lines from erect and pitched planes. Perception, 25, 831-852. Mermelstein, R., Banks, W., & Prinzmetal, W. (1979). Figural goodness effects in perception and memory. Perception & Psychophysics, 26, 472-480. Muir, D., & Over, R. (1970). Tilt aftereffect in central and peripheral vision. Journal of Experimental Psychology, 85, 165-170. Pressey, A. (1971). An extension of assimilation theory to illusions of size, area, and direction. Perception & Psychophysics, 9, 172-176. Pressey, A. W., Butchard, N., & Scrivner, L. (1971). Assimilation theory and the Ponzo illusion: Quantitative predictions. Canadian Journal of Psychology, 25, 486-497. Pressey, A. W., & Epp, D. (1992). Spatial attention in Ponzo-like patterns. Perception & Psychophysics, 52, 211-221. Pressey, A. W., & Sweeney, O. (1972). Acute angles and the Poggendorff illusion. Quarterly Journal of Experimental Psychology, 24, 169-174. Prinzmetal, W., Shimamura, A. P., & Mikolinski, M. (in press). The Ponzo illusion and the tilt-constancy theory. Perception & Psychophysics. Robertson, L. C. (1995). Covert orienting biases in scene-based reference frames: Orientation priming and visual field differences. Journal of Experimental Psychology: Human Perception and Performance, 21, 707-718. Rock, I. (1974). The perception of disoriented figures. Scientific American, 230, 78-86. Sauvan, X. M., & Peterhaus, E. (1995). Neural integration of visual information and direction of gravity in prestriate cortex of the alert monkey. In T. Mergner & F. Hlavacka (Eds.), Multisensory control of posture (pp. 43-49). New York: Plenum Press. Shimamura, A. P., & Prinzmetal, W. (2000). The mystery spot illusion and its relation to other visual illusions. Psychological Science, 10, 501-507. Singer, G., Purcell, A. T., & Austin, M. (1970). The effect of structure and degree of tilt on the tilted room illusion. Perception & Psychophysics, 7, 250-252. Spivey-Knowlton, M., & Bridgeman, B. (1993). Spatial context affects the Poggendorff illusion. Perception & Psychophysics, 53, Mil-414. Stoper, A. E. (1998). Height and extent: Two kinds of size perception. In E. Winograd, R. Fivush, & W. Hirst (Eds.), Ecological approaches to cognition (pp. 97-121). Hillsdale, NJ: Erlbaum. Stoper, A. E., & Cohen, M. M. (1986). Judgments of eye level in light and in darkness. Perception & Psychophysics, 40, 311-316. Templeton, W. B. (1973). The role of gravitational cues in the judgment of visual orientation. Perception & Psychophysics, 14, 451-457. 217 Tyler, C. W., & Nakayama, K. (1984). Size interactions in the perception of orientation. In L. Spillman & B. R. Wooten (Eds.), Sensory experience, adaptation, and perception: Festschrift for Ivo Kohler (pp. 529546). Hillsdale, NJ: Erlbaum. Velinsky, S. (1925). Explication physiologique de l'illusion der Poggendorff [Physiological explanation of the Poggendorff illusion]. Annie Psychologique, 107-116. Wallace, G. K., & Moulden, B. (1973). The effect of body tilt on the Zollner illusion. Quarterly Journal of Experimental Psychology, 25, 10-21. Ward, L. M., & Coren, S. (1976). The effect of optically induced blur on the magnitude of the Mueller-Lyer illusion. Bulletin of the Psychonomic Society, 7, 483-484. Weintraub, D. J., & Krantz, D. H. (1971). Poggendorff illusion: Amputations, rotations, and other perturbations. Perception & Psychophysics, 10, 257-263. Weintraub, D., Krantz, D., & Olson, T. (1980). The Poggendorff illusion: Consider all angles. Journal of Experimental Psychology, 6, 718-725. Wenderoth, P. M. (1974). The distinction between the rod-and-frame illusion and the rod-and-frame test. Perception, 3, 205-212. Wenderoth, P., & Johnstone, S. (1988). The mechanism of the direct and indirect tilt illusions. Vision Research, 28, 301-312. Wilson, A. E., & Pressey, A. W. (1976). The role of apparent distance in the Poggendorff illusion. Perception & Psychophysics, 20, 309-316. Witkin, H. A., & Asch, S. E. (1948a). Studies in space orientation: III. Perception of upright in absence of a visual field. Journal of Experimental Psychology, 38, 603-614. Witkin, H. A., & Asch, S. E. (1948b). Studies in space orientation: IV. Further experiments of the upright with displaced visual fields. Journal of Experimental Psychology, 38, 762-782. Wolfe, J., & Held, R. (1982). Gravity and the tilt aftereffect. Vision Research, 22, 1075-1078. Yin, R. (1969). Looking at upside-down faces. Journal of Experimental Psychology, 81, 141-145. Zoccolotti, P., Antonucci, G., & Spinelli, D. (1993). The gap between rod and frame influences the rod-and-frame effect with small and large inducing displays. Perception & Psychophysics, 54, 14-19. Received April 20, 1999 Revision received May 8, 2000 Accepted May 12, 2000