Visual and Auditory Influence on Perceptual Stability in Visual Competition
Identifieur interne : 002076 ( Istex/Corpus ); précédent : 002075; suivant : 002077Visual and Auditory Influence on Perceptual Stability in Visual Competition
Auteurs : Katsumi Watanabe ; Kohske TakahashiSource :
- Seeing and Perceiving [ 1878-4755 ] ; 2011.
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DOI: 10.1163/187847511X588809
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<p>Seeing and Perceiving 24 (2011) 545–564 brill.nl/sp Visual and Auditory Influence on Perceptual Stability in Visual Competition ∗ Kohske Takahashi 1 , 2 , ∗∗ and Katsumi Watanabe 1 , 3 1 Research Center for Advanced Science and Technology, The University of Tokyo, 4-6-1 Komaba, Meguro-ku 153-8904, Tokyo, Japan 2 Japan Society for the Promotion of Science, 5-3-1 Koujimachi, Chiyoda-ku, Tokyo 102-8471, Japan 3 Japan Science and Technology Agency, 4-1-8, Honcho, Kawaguchi-shi, Saitama 332-0012, Japan Received 30 January 2011; accepted 28 June 2011 Abstract In visual competition, the perception of ambiguous visual patterns changes spontaneously. Although the process causing this perceptual alternation remains unclear, recent evidence suggests various types of non- visual influences in resolving visual ambiguity. In the present study, we investigated cross-modal modulation of a transient stimulus on visual perceptual stability (i.e., alternation frequency). Participants observed an ambiguous visual figure and reported their perceptual alternations. Concurrently, we presented visual and auditory transient events. The results revealed that the auditory as well as visual transient events destabilize the current perception (i.e., they increase alternation frequency) around 0.5–1.5 s after the event. In addition, the magnitudes of auditory and visual effects were comparable and positively correlated within participants. These results suggest that the visual perceptual stability can be under the influence of processes that are shared by different senses. © Koninklijke Brill NV, Leiden, 2011 Keywords Visual competition, perceptual stability, vision, audition, cross-modal interaction 1. Introduction Perception of competitive visual input changes from one to the other (visual com- petition; e.g., Blake and Logothetis, 2002; Kim and Blake, 2005; Leopold and Logothetis, 1999; Sterzer and Kleinschmidt, 2007; Sterzer et al. , 2009; Tong et * This article is part of the Multisensorial Perception Collection, guest edited by S. Wuerger, D. Alais and M. Gondan. ** To whom correspondence should be addressed. E-mail: ktakahashi@fennel.rcast.u-tokyo.ac.jp © Koninklijke Brill NV, Leiden, 2011 DOI:10.1163/187847511X588809</p>
<p>546 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 al. , 2006). When different images are projected independently to the left and right eyes, they cannot simultaneously arise in visual awareness. In fact, the image aris- ing in visual awareness spontaneously moves between the images in both the left and right eyes (binocular rivalry). Another form of visual competition is known as stimulus rivalry (e.g., Necker cube). When the number of possible interpretations of a visual figure is more than one, the interpretation arising in visual awareness spontaneously switches from one to the other. Visual competition is characterized by two main factors — dominant percept and perceptual stability. Dominant percept refers to which of the possible interpre- tations arises in visual awareness, and perceptual stability refers to how frequently perception changes. Several researches have revealed that dominant percept and perceptual stability are susceptible to modulations by various factors such as at- tention (Chong and Blake, 2006; Chong et al. , 2005; Khoe et al. , 2008; Meng and Tong, 2004; Mitchell et al. , 2004; Paffen et al. , 2006; Tsal and Kolbet, 1985), in- tention (Kornmeier et al. , 2009; Suzuki and Peterson, 2000; Toppino, 2003), action (Maruya et al. , 2007), task-irrelevant visual stimulation (Freeman and Driver, 2006; Kanai et al. , 2005; Paffen et al. , 2005), stimulus configuration (Blake et al. , 2003; Freeman and Driver, 2006; Ilg et al. , 2008; Leopold et al. , 2002), perceptual history (Brascamp et al. , 2008; Knapen et al. , 2009; Maier et al. , 2003; Naber et al. , 2010; for review, Pearson and Brascamp, 2008), eye movement and retinal image shift (van Dam and van Ee, 2005, 2006a, 2006b), and transcranial magnetic stimulation to visual areas (Brascamp et al. , 2010; Pearson et al. , 2007). These results suggest that visual competition reflects not only low-level competition in sensory specific processes but also perceptual and cognitive modulation on visual processes. Besides these visual or attentional modulations on visual competition, several researches have shown cross-modal effects in resolving visual ambiguity; sensory inputs other than vision can influence visual competition, especially dominant inter- pretation. For example, sound alters the interpretation of ambiguous visual events (e.g., stream-bounce illusion, Sekuler et al. , 1997; Watanabe and Shimojo, 2001), and touching haptic objects biases the interpretation of the Necker cube or the struc- ture from motion stimulus (Ando and Ashida, 2003; Blake et al. , 2004; Bruno et al. , 2007; James and Blake, 2004). Recently, van Ee et al. (2009) demonstrated that visual inputs that shared temporal configurations with auditory or tactile inputs tended to be dominant percepts in resolving binocular rivalry. These results imply that dominant percepts are intrinsically biased to maintain cross-modal congruen- cies among inputs into multiple sensory modalities. As compared to the dominant percept, cross-modal modulation on perceptual sta- bility is less understood. Bruno et al. (2007) addressed this issue by revealing that the alternation frequency from consistent to inconsistent percepts decreased after haptic inputs. In their study, observers reported the percept of a visual Necker cube while touching a haptic cube. After the transition from stationary touch to moving touch, a switch from consistent to inconsistent visual percepts was suppressed. In other words, perceptual stability increased when visual and tactile inputs were con-</p>
<p>K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 547 gruent. More recently, Takahashi and Watanabe (2010) reported that auditory events could influence perceptual stability even without cross-modal congruencies using the learning paradigm. They found that after exposure to temporally synchronized audiovisual events, alternation frequencies decreased in the absence of auditory events. They also found that alternation frequencies tended to increase immediately after auditory transient events. The latter finding would be related to visually in- duced perceptual alternation (visual IPA; Kanai et al. , 2005). These results imply that inputs into sensory modalities other than vision can both increase and decrease visual perceptual stability. Although non-visual influences on visual perceptual stability have been reported, they have not been compared directly with visual influence. Therefore, it is still unclear whether visual and auditory/tactile transients influence visual perceptual stability in a similar manner. In this study, we examined whether task-irrelevant visual and auditory transient events would induce or suppress perceptual alternation while experiencing stimulus rivalry and, if so, whether the effects of these visual and auditory events would be comparable. If visual and auditory transients affect visual alternation in a similar fashion, it would imply that visual stability could be under the influence of cross-modal interaction. 2. Experiment 1 2.1. Methods 2.1.1. Apparatus and Stimuli The visual stimulus was an ambiguous apparent motion (quartet dot) that consisted of two blue dots (0.36 ◦ , 17.0 cd/m 2 ) against a gray background (15.7 cd/m 2 ) on a 21-inch CRT monitor (85 Hz; viewing distance: 57 cm) (Fig. 1(a)). The two dots alternately appeared on the top-left and bottom-right vertices or bottom-left and top-right vertices of an imaginary square (2 . 7 ◦ × 2 . 7 ◦ ) placed in the center of the Figure 1. (a) Experimental procedures. An ambiguous visual figure (apparent motion) was presented concurrently with transient events (denoted using gray circles). The transient events were repeatedly presented during 240 s session with the random temporal intervals sampled from 1-s-bin discrete uni- form distribution ranging over the interval 4–12 s. Participants traced the perceived motion direction of the ambiguous figure and reported the perceptual alternation (denoted using black squares). The elapsed time of the perceptual alternation from the preceding transient event was defined as tempo- ral lag. (b) Schematic illustration of the transient events. Onset of the transient events was always synchronized with the timing of the change of dot position.</p>
<p>548 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 screen. The duration of each display was 500 ms. The visual transient event was a white (79.2 cd/m 2 ) flash of the background lasting 105 ms. The auditory transient event was a beep sound lasting 100 ms that was presented through headphones. 2.2. Procedure Twenty-eight undergraduate students participated in Experiment 1. A session began when a participant pressed the enter key. The apparent motion stimulus appeared af- ter a blank period that lasted for 1 s. The participants traced the perceived direction of the apparent motion (horizontal or vertical) by pressing or releasing the space bar. A number of transient events were presented intermittently during a 240 s session. The temporal intervals between the onsets of two successive transient events were randomly sampled from a 1 s bin discrete uniform distribution ranging from 4 to 12 s (Fig. 1(a)). The onset of the transient events was synchronized with the update of apparent motion display (Fig. 1(b)). The participants were explicitly instructed to ignore the transient events. We conducted two conditions — visual and auditory – as per the sensory modality of the transient events. These conditions were sepa- rately conducted in a 240 s session and were repeated twice in random order. Before starting the experiment, we confirmed that the participants experienced perceptual alternation with the ambiguous motion display, after which they performed one 30 s session for practice. 2.3. Results and Discussion The overall alternation frequency, which was defined as the total number of per- ceptual alternations in each condition (480 s), did not differ between the visual and auditory conditions (visual: 33.1, auditory: 31.2 times; t ( 27 ) = 0 . 36, p = 0 . 72), suggesting that the overall alternation frequency was independent of the sensory modality of the task-irrelevant transient events. The alternation frequency showed large individual differences ranging from 0–135 times/480 s (Leopold and Logo- thetis, 1999). The large individual difference of the overall alternation frequency might originate from the following two factors: (1) the variation of the rate of per- ceptual alternation and (2) the variation of the criterion for perceptual alternation. In this study, we examined the timing of perceptual alternation averaged for each par- ticipant in each condition so as to assess the effect magnitudes of transient events. The averaged timing would inevitably be unreliable when the alternation frequency was extremely low. Therefore, when the alternation frequency was less than 5 in at least one condition, we excluded the participants from the following analyses (7 participants). Note that the exclusion had little influence on the distribution anal- yses (Fig. 2), since the total number of perceptual alternations of these participants was so small that they could be ignored. To examine the timing-selective modulation of transient events on visual per- ceptual stability, we first calculated the elapsed time of each perceptual alternation from the preceding transient events, which we defined as ‘temporal lag’ (Fig. 1(a); see also Kanai et al. , 2005; Takahashi and Watanabe, 2010), and then examined the</p>
<p>K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 549 Figure 2. Normalized frequency (frequency per 1 s) of each 0.5 s bin (a), cumulative distribution func- tions (b), and differences of distribution function (observed–resampled) (c) of temporal lag. Temporal lag was defined as the elapsed time of each perceptual alternation from the preceding transient event. The solid and dotted lines in (b) show the temporal lags calculated using real and resampled transient events, respectively.</p>
<p>550 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 distributions of the temporal lags (see Note 1). For example, if a perceptual alterna- tion took place 9.5 s after the session began and if the preceding transient occurred 6 s after the session began, the temporal lag of the perceptual alternation is 3.5 s. Figure 2(a) shows the normalized frequency of the temporal lag (frequency per 1 s) of each 0.5 s bin. If a transient event immediately induces perceptual alternation, the frequency of small temporal lags would be higher than large temporal lags. On the other hand, if transient events suppress perceptual alternation for the sub- sequent period, the frequency of small temporal lags would be lower. Otherwise, if the transient event has no timing-selective influence on perceptual alternation, the distribution of temporal lag would have no peak or gradient but would be uniform. Inspection of the data suggested that the frequency of temporal lag was relatively high around 0.5–1.5 s compared to the other period. In other words, perceptual al- ternation frequently occurred around 0.5–1.5 s after the transient events took place. To ensure that the non-uniform distribution of the temporal lag was truly due to the transient event, we simulated the experimental sessions using the timings of the observed perceptual alternations and those of the newly sampled simulated tran- sient events discarding the actually presented transient event. We also calculated the ‘resampled temporal lag’ in the same way as the temporal lag. The resampled tem- poral lag gives the approximation of the theoretical distribution of the temporal lag under the assumption that the timing of perceptual alternation is independent from those of the transient events. The cumulative distribution functions of the observed and resampled temporal lags differed significantly in both conditions (Fig. 2(b), two-sample Kolmogorov–Smirnov test, D s > 0 . 16, p s < 10 − 14 ). The difference was elevated around 0.5–2 s after the transient event occurred (Fig. 2(c)); the effect period was similar to those in visual IPA (Kanai et al. , 2005). Analyses of the distributions of temporal lag clearly showed that the transient events induced timing-specific destabilization. To evaluate the magnitude of the destabilizing effect of the individuals, we calculated the arithmetic mean of tem- poral lags for each participant in each condition (mean temporal lag; Fig. 3(a)). If perceptual alternation frequently takes place immediately after the transient event, the mean of the distribution of the temporal lag shifts becomes smaller, which im- plies a stronger destabilizing effect. Therefore, the mean temporal lag is one of the indices that represent the magnitude of the destabilizing effect (see Note 2). The mean temporal lags were significantly smaller than chance in both conditions (one- sample two-tailed t -test, visual: t ( 20 ) = 3 . 46, p < 0 . 01; auditory: t ( 20 ) = 2 . 93, p < 0 . 01). Furthermore, the mean temporal lag did not significantly differ between the visual and auditory conditions ( t ( 20 ) = 1 . 90, p = 0 . 07) (see Note 3). Taken to- gether, these results indicated that both auditory and visual transients immediately induced perceptual alternation at the comparable magnitudes. Next, we addressed the individual differences of the magnitudes of the destabi- lizing effect and the overall alternation frequency using within-subject correlation analysis. A positive correlation was found between the visual and auditory condi- tions with regard to the mean temporal lag (Fig. 3(b), r = 0 . 59, p < 0 . 01) as well as</p>
<p>K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 551 Figure 3. (a) Mean temporal lags. The horizontal line indicates the chance levels (4.41 s). The error bars indicate the standard errors of the means. (b)–(e) Within-subject correlations. Each point repre- sents an individual participant. Solid lines show the linear regression. (b) Mean temporal lags in the visual and auditory conditions. (c) Overall alternation frequency in the visual and auditory conditions. (d and e) Mean temporal lags and the overall alternation frequency in the visual (d) and auditory (e) conditions.</p>
<p>552 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 the overall alternation frequency (Fig. 3(c), r = 0 . 64, p < 0 . 01). These results indi- cate that the magnitude of the destabilizing effects of transient events and the overall alternation frequency varied among individuals, but they were relatively constant between the modalities of the transient event. However, the mean temporal lag and the alternation frequency did not correlate in both the visual (Fig. 3(d), r = 0 . 02, p = 0 . 94) and auditory (Fig. 3(e), r = − 0 . 12, p = 0 . 62) conditions. Perhaps, the overall alternation frequency would reflect the intrinsic visual perceptual stability in observing ambiguous figures, and the destabilizing effect would reflect the suscep- tibility of visual perceptual stability to the external transient events. Therefore, the susceptibility to the external event might not be related to the intrinsic perceptual stability. Finally, we found a directional bias for the quartet dot stimulus (Hock et al. , 1996). The participants preferably perceived vertical motion over horizontal motion (73% vs. 27%). However, within-correlation analyses showed that the magnitude of directional bias was not related to the strength of destabilizing effect (visual: r = − 0 . 12, p = 0 . 62; auditory: r = 0 . 02, p = 0 . 92). Moreover, the mean temporal lag did not differ between two types of perceptual alternation ( F ( 1 , 20 ) = 1 . 60, p = 0 . 22) — vertical-to-horizontal and horizontal-to-vertical — thus, the destabilizing effect would reflect the change of stability irrespective of the dominant percept. In sum, we replicated visual IPA (Kanai et al. , 2005) and also found a similar effect using auditory transient events. In addition, the magnitudes of the visual and auditory destabilizing effects were correlated, but they did not correlate with the overall alternation frequencies. 3. Experiment 2 Although the results of Experiment 1 showed that visual and auditory transient events influenced visual perceptual stability, it was still unclear whether the alterna- tion frequency increased immediately after the transient events or decreased while the transient events were absent. For example, Takahashi and Watanabe (2010) found both a timing-selective increase and decrease of alternation frequency due to external events. In Experiment 2, therefore, we compared the alternation frequency between the situations with and without transient events. 3.1. Methods The apparatus, stimuli, and procedures were identical to those in Experiment 1, except for the following points. In Experiment 2, we presented neither visual nor auditory transient events in one condition (no-transient condition) in addition to the visual and auditory conditions. Fourteen participants were newly recruited for this experiment. The aspect ratio of the quartet dot stimulus was modified (2.7 ◦ width × 3 . 4 ◦ height) in order to reduce the directional bias.</p>
<p>K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 553 3.2. Results and Discussion The overall alternation frequency did not differ among conditions (visual: 34.6, au- ditory: 41.4 and no-transient: 43.2 times s; one-way repeated measures ANOVA, F ( 2 , 26 ) = 0 . 37, p = 0 . 69). The overall frequency showed large individual differ- ences ranging from 1–169 times/480 s. We excluded participants on the same basis of alternation frequencies as in Experiment 1 (3 participants). The effects of the transient events in the visual and auditory conditions were similar to those in Experiment 1. The normalized frequency of temporal lags of each 0.5 s bin was relatively high at around 0.5–1.5 s (Fig. 4(a)). The cumula- tive distributions of the observed and resampled temporal lags differed significantly in both conditions (Fig. 4(b), all D s > 0 . 20, p s < 10 − 15 ), and the effect periods (Fig. 4(c)) were also similar to those in Experiment 1. The quantitative evaluation using mean temporal lags for each participant (Fig. 5(a)) showed that the mean temporal lags were significantly smaller than chance (one-sample two-tailed t -test, visual: t ( 10 ) = 2 . 57, p < 0 . 05; auditory: t ( 10 ) = 2 . 52, p < 0 . 05), and also that the magnitude of the destabilization effect did not differ between the visual and audi- tory conditions ( t ( 10 ) = 0 . 32, p = 0 . 76). A series of correlation analyses revealed that the mean temporal lag (Fig. 5(b), r = 0 . 77, p < 0 . 01) as well as the overall al- ternation frequency (Fig. 5(c), r = 0 . 66, p < 0 . 05) positively correlated between the auditory and visual conditions within individuals, while the mean temporal lag and the alternation frequency did not correlate in either the visual (Fig. 5(d), r = 0 . 00, p = 0 . 99) or auditory conditions (Fig. 5(e), r = 0 . 15, p = 0 . 65) (see Note 4). Thus, we replicated the results of Experiment 1, suggesting that visual and auditory modulation on alternation frequency is a robust effect, at least for the participants whose overall alternation frequency was not too small. Finally, we compared the frequency of perceptual alternation after the transient events with the frequency in the no-transient condition. We calculated the ‘default’ normalized frequency of temporal lag, which indicates how frequently perceptual alternation can take place without the transient events, by dividing the overall alter- nation frequency in the no-transient condition by the duration of session (480 s) (the horizontal lines in each panel of Fig. 4(a)) (see Note 5). Visual inspection showed that the alternation frequencies for 0.5–1.5 s after the transient events were much higher than that in the no-transient condition, while the alternation frequencies for the subsequent period were slightly lower than that in the no-transient condition. To evaluate this, we compared the normalized frequencies of temporal lag for 0.5– 1.5 s after the transient events (effective period), those for the subsequent 1.5–12 s period (non-effective period), and those in the no-transient condition (no-transient period). A one-way repeated measures ANOVA revealed the significant main effect of period ( F ( 2 , 20 ) = 5 . 03, p < 0 . 05). The normalized frequencies for the effective period were significantly higher than those in the no-transient period ( t ( 10 ) = 2 . 29, p < 0 . 05, correction for multiple comparison using Ryan’s method) and the non- effective period ( t ( 10 ) = 3 . 05, p < 0 . 01). The difference between non-effective and no-transient periods did not reach significance ( t ( 10 ) = 0 . 75, p = 0 . 46). These</p>
<p>554 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 Figure 4. Normalized frequency (frequency per 1 s) of each 0.5 s bin (a), cumulative distribution functions (b), and differences of distribution function (observed–resampled) (c) of temporal lag. For details, see Fig. 2. The horizontal line in (a) indicates the baseline alternation frequency that was calculated using no-transient condition.</p>
<p>K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 555 Figure 5. (a) Mean temporal lags. (b)–(e) Within-subject correlations. For details, see Fig. 3. results showed that perceptual alternation frequently takes place immediately af- ter the transient events as compared to the ‘default’ situation wherein the transient events were absent throughout the session.</p>
<p>556 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 In Experiment 2, the directional bias was reduced by modifying the aspect ratio (46% for vertical motion and 54% for horizontal motion). Furthermore, as well as the Experiment 1, the results of within-subject correlation between the directional bias and temporal lag (visual: r = 0 . 57, p = 0 . 07; auditory: r = 0 . 06, p = 0 . 85) and the temporal lag separated into two types of alternation ( F ( 1 , 10 ) = 0 . 13, p = 0 . 72) confirmed that the destabilization effect would not be related to the dominant percept. 4. General Discussion In the present study, we investigated how task-irrelevant visual and auditory tran- sient stimulation would influence perceptual stability, i.e., alternation frequency, while observing stimulus rivalry. The results clearly demonstrated the timing- selective modulation of both visual and auditory transient events on visual percep- tual stability; the alternation frequency for ambiguous visual figures was elevated around 0.5–1.5 s after the transient events took place, and the magnitude of the destabilizing effects were comparable and correlated between visual and auditory transient events. We also ensured that the transient events increased alternation fre- quency for subsequent periods compared to the baseline frequency without transient events. In the context of cross-modal influence on visual competition, several researches have reported that auditory and tactile stimuli that had cross-modal congruencies with visual stimuli could bias dominant percepts (Ando and Ashida, 2003; Blake et al. , 2004; Bruno et al. , 2007; James and Blake, 2004; van Ee et al. , 2009). These biases could lead to the distortion of the alternation frequency; when the auditory or tactile inputs were congruent with the visual inputs, perceptual alternation to in- congruent percepts would be suppressed (e.g., Bruno et al. , 2007). In such cases, the change in perceptual stability would not be essential but would be a subsidiary effect. However, the timing-selective destabilizing effect observed in the present study would not be a subsidiary effect of the bias of dominant percepts since the transient events had no cross-modal congruencies with the bistable visual pattern. Takahashi and Watanabe (2010) also showed that auditory events both increase and decrease perceptual stability even in the absence of the bias toward dominant per- cepts. Therefore, sound might influence perceptual stability but not as the result of the bias to dominant percept. Visual transients are known to decrease perceptual stability and induce percep- tual alternation for subsequent periods (Kanai et al. , 2005). In our study, both visual and auditory transient events increased, not suppressed, the perceptual alternation, and the effect period was around 0.5–1.5 s after the transient events (Figs 2 and 4). Since there was a 500 ms interval between visual stimuli in the present experiment, the effects of transients on the motion direction percept could not be manifested un- til the following visual stimuli was presented after the transient event. By subtract- ing 0.5 s from the elapsed time from the transient events to perceptual alternation,</p>
<p>K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 557 the influence on perceptual stability would immediately begin after the transients. The time courses of the effects between visual and auditory transient events were quite similar, and resembled those in Kanai et al. (2005, see their Fig. 2). Further- more, quantitative analyses revealed that the effect magnitudes, i.e., mean temporal lag, were comparable and showed within-subject correlation between visual and auditory transient events (Figs 3(a), (b), 5(a) and (b)). These results imply that the influence of visual and auditory transients on perceptual stability might be mediated by similar mechanisms, at least for those individuals whose alternation frequency is not too small. Thus, our findings would extend the influence of transient events on perceptual stability into cross-modal domains; the auditory as well as visual tran- sient events reduced the visual perceptual stability of the current percept regardless of the content of the percept. In Experiment 1, we found a significant directional bias toward vertical motion (Hock et al. , 1996). Therefore, the effects of transient events could be to bias the current percept to the dominant one. However, the correlation analyses indicated that directional bias and the magnitudes of destabilizing effect were not related. We also found the destabilizing effect even in Experiment 2, where the directional bias was eliminated, and again directional bias and the strength of destabilizing effect did not correlate. As well, the temporal lag for two types of alternation — horizontal-to-vertical and vertical-to-horizontal — was comparable. Another ac- count might relate to the decision criteria. In binocular rivalry and some types of stimulus rivalry (e.g., structure from motion), observers sometimes experience the two competitive states as partially intermixed and being ambiguous (Blake and Logothetis, 2002). In such cases, change of decision criteria would distort temporal characteristics of perceptual alternation. However, in the quartet-dot stim- ulus, the ambiguous state was less conspicuous, and therefore we think that the change of decision criteria would be unlikely to explain the destabilization effect. These observations suggest that the destabilizing effect may reflect processes for visual perceptual stability, irrespective of the dominant percept or the decision cri- teria. How do visual and auditory transients influence perceptual stability and induce perceptual alternations? Visual and auditory transients might induce immediate eye movement, leading to perceptual alternation. Indeed, eye-movements were asso- ciated with the perceptual alternation in binocular rivalry (van Dam and van Ee, 2006a, 2006b). However, although the effect of eye-movement in stimulus rivalry is still debated, several researchers reported that perceptual alternation in stimulus rivalry is not associated with eye movements including saccades, micro-saccades, or blink, as opposed to binocular rivalry (Ito et al. , 2003; van Dam and van Ee, 2005, 2006a, 2006b). In addition, Kanai et al. (2005) demonstrated the spatial se- lectivity of the transient effect and suggested that eye movement would not be the main reason for the transient-induced alternation. Taken together, we favor the idea that eye movements do not play a key role in the destabilization effect. However,</p>
<p>558 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 since we did not directly measure eye-movements during the session in the present study, further research is required to examine this issue. Attention may be responsible for both visual and auditory transient-induced alternation. The task-irrelevant transient events inevitably attract visual attention (Posner and Cohen, 1984), which would lead the focused attention away and re- orient it to the bistable figure. As several findings support the involvement of atten- tional system in perceptual alternation (Chong and Blake, 2006; Chong et al. , 2005; Georgiades and Harris, 1997; Khoe et al. , 2008; Meng and Tong, 2004; Mitchell et al. , 2004; Paffen et al. , 2006; Tsal and Kolbet, 1985), Kanai et al. (2005) argued that visually induced perceptual alternation would be mediated by the exogenous shift of visual attention. Perhaps this account can be extended into cross-modal domain. As for cross-modal attention, several studies have revealed that auditory in- puts exogenously modulate visual attention (Mazza et al. , 2007; Spence and Driver, 1997). Moreover, neuroimaging studies suggest that perceptual alternation would be related to the activation and fluctuation of the parietal cortex (Britz et al. , 2009, 2011; Kanai et al. , 2010), and the parietal cortex mediates attention in non-visual modalities (Shomstein and Yantis, 2006). In the present study, we showed that the influences of visual and auditory transient events were similar both qualitatively and quantitatively as well as positively correlated within-subject, implying that similar mechanisms might underlie these effects. Such evidence suggests that both visual and auditory transients induce the exogenous shift of visual attention that is medi- ated by the parietal cortex, leading to destabilization in visual competition. In the case of auditory modulation, relatively higher-level, top–down modulation might serve to induce the visual attentional shift. Meanwhile, recent findings suggest that the sensory processes of visual and other modalities are not completely independent (Alais et al. , 2010; Driver and Noesselt, 2008; Shams and Kim, 2010; Shimojo and Shams, 2001). In fact, auditory transient events influence early visual sensory processes. For example, when a single flash of light is accompanied by double auditory transient stimulation, people experience the flash twice (Shams et al. , 2000, 2002). A large body of research suggested that the auditory transient event could modulate the primary visual areas (Mishra et al. , 2007, 2008; Shams et al. , 2005; Watkins et al. , 2006, 2007). Therefore, the auditory as well as visual transients might directly induce fluctuation in the visual area, lead- ing to destabilization. Note that Mishra et al. (2010) recently demonstrated that the auditory influences on the visual cortex involved the attentional process. The cross- modal interaction mediated by the attentional process in the parietal cortex and that by the low-level direct connection might interactively induce destabilization. Thus, although current evidence would not be sufficient to identify the underlying neu- ral mechanisms of visual and auditory induced perceptual alternation, visual and auditory transients might influence visual sensory processes relevant to perceptual stability both directly and via the exogenous shift of attention. Observing that the visual and auditory transient events induced destabilization in a similar manner, the destabilization would not concern the sensory modality at</p>
<p>K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 559 which the transient events arise. Rather, the saliency of the transient events might matter. Since we presented the transient events with an abrupt onset and offset, these events had the lowest temporal uncertainty and hence, the highest saliency. However, increasing temporal uncertainty weakens the power of transients (e.g., Van der Burg et al. , 2010). Therefore, to determine if any type of event is sufficient or whether or not a higher saliency is necessary to induce destabilization, it would be worthwhile to examine whether the auditory and visual events with higher tem- poral uncertainty could induce destabilization at similar magnitudes as the transient events. The within-subject correlation analyses revealed that the magnitudes of the destabilizing effect of the visual and auditory transient events positively correlated (Figs 3(b) and 5(b)). Similarly, the overall alternation frequency also positively cor- related between the visual and auditory conditions (Figs 3(c) and 5(c)). Meanwhile, the magnitudes of the destabilizing effect and overall alternation frequency did not correlate (Figs 3(d), (e), 5(d) and (e)). These results imply that spontaneous and induced alternation may reflect distinct mechanisms. The spontaneous alternation rate might be determined mainly through bottom-up processes, such as intrinsic neural oscillators (Carter and Pettigrew, 2003) that are modality-specific (Hupé et al. , 2008; Pressnitzer and Hupé, 2006) and insensitive to external events. Indeed, Kanai et al. (2010) reported that the structure of the parietal cortex could explain the individual differences in the rate of spontaneous alternation. On the other hand, the top-down process, which is modality-unspecific and tuned into external tran- sient events, would timing-selectively modulate visual perceptual stability. In the present study, we examined the magnitudes of the effects for participants whose alternation frequencies were not too small in order to ensure statistical reliability. Therefore, the positive correlation between the effects of visual and auditory tran- sients might be limited to those participants, and it is still unclear whether the same results would be observed for the individuals showing very much smaller alterna- tion frequencies. In summary, the present study demonstrates that both visual and auditory tran- sient events modulate visual perceptual stability. The exogenous influence of tran- sient events on perceptual stability may be shared among different sensory modal- ities. For example, the question of whether visual and auditory transient events influence auditory rivalry (e.g., auditory streaming, Bregman and Campbell, 1971) still remains answered. Some studies suggest that the activity of the intra-parietal sulcus was related to auditory rivalry (Cusack, 2005), implying that transient events might influence auditory rivalry as well as visual competition through the activa- tion or fluctuation in the parietal cortex. On the other hand, Pressnitzer and Hupé (2006) showed that within-subject correlation between visual and auditory rivalry was weak, implying that different mechanisms underlie the auditory and visual competition. Further research examining the rivalry in diverse sensory modalities will shed light on perceptual stability in a cross-modal scheme.</p>
<p>560 K. Takahashi, K. Watanabe / Seeing and Perceiving 24 (2011) 545–564 Acknowledgements This work was supported by the Japan Science and Technology Agency and the Japan Society for the Promotion of Science. We would like to thank Shinsuke Shi- mojo and Shohei Hidaka for their helpful comments and technical advice. Notes 1. The distribution of the temporal lag is not related to the overall alternation fre- quency. 2. The advantages of using the mean temporal lag as the index of effect magnitude is that the arithmetic mean is distribution-free and that we can calculate the chance level assuming that transient events have no timing-specific effect on perceptual alternation. 3. If the timings of perceptual alternation are independent from those of the tran- sient events, the probability density function of the temporal lag is as follows: P ( lag ) = 1 72 12 ∑ δ = 4 H (δ − lag ), where H is a Heaviside function (see also Takahashi and Watanabe, 2010, pp. 3–4). The expected value (i.e., mean) of the density function is 4.41 s, which is determined only by the distribution of the temporal intervals of the transient event, independent of the overall alternation frequency. The overall alternation frequency can be relevant to the sample size. The more frequent alternation (i.e., the larger sample size) leads to the smaller variance in estimating the mean of the temporal lag, while the sample size does not distort the estimated mean of the distribution. 4. We noted that one participant in Experiment 2 was a possible outlier (Fig. 4(b) and (d)), so we performed statistical tests excluding the participant. The results were almost unchanged except for that correlation between temporal lag of vi- sual and auditory condition (Fig. 4(b)) was not significant (but still positive, r = 0 . 36, p = 0 . 29). 5. If the transient events had no effect, the normalized frequency of each bin must be identical to the default normalized frequency. The higher (lower) normal- ized frequency than default indicates that perceptual alternation more (less) frequently took place in the bin. References Alais, D., Newell, F. N. and Mamassian, P. (2010). Multisensory processing in review: from physiol- ogy to behavior, Seeing and Perceiving 23 , 3–38. Ando, H. and Ashida, H. (2003). Touch can influence visual depth reversal of the Necker cube, Per- ception 32 , 97.</p>
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