85 Study IV. A new paradigm for measuring the ERP correlates of auditory multistable
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General discussion
The main goal of the studies presented in this thesis was to investigate the percept-inducing and percept-stabilizing cues and the ERP correlates of the perceptual processes of auditory stream segregation. The studies reported here utilized behavioral and electrophysiological measurements for better understanding auditory stream segregation.
Continuous measurements during bi-/multistable perception permit one to investigate various aspects of stream segregation while eliminating the effects originating from physical stimulus changes. Therefore, multistable stimulation was used in all four studies while the participant’s perception of the stimuli was continuously recorded. Our results showed that similarly to other cues based on spectral similarity/separation, amplitude modulation can both induce and stabilize auditory stream segregation, and it interacts with carrier frequency difference (Study I). We demonstrated that some cues based on spectro-temporal regularities, such as a melody, may also induce stream segregation (Study II). Studies on the neural correlates of auditory stream segregation (Study III and IV) demonstrated that the current perceptual interpretation influences sound processing both at the level of feature extraction and sound evaluation.
Percept-inducing and stabilizing cues
Whereas most of the studies on auditory stream segregation used cues based on spectral differences between the sounds (Hartmann & Johnson, 1991) there is much evidence showing that the auditory system also utilizes other types of cues (for reviews, see Moore &
Gockel, 2002; Moore & Gockel, 2012). Grimault et al. (2002) found that differences between the modulation frequency of two sets of sounds can evoke auditory stream segregation. Study I investigated the effects of amplitude modulation in a bistable auditory streaming paradigm together with carrier-frequency differences and differences in the perceived location. Based on the results, amplitude modulation difference not only stabilized streams but also induced
98 stream segregation, and the effect of amplitude modulation difference was partly overlapping with that of the carrier frequency difference and location difference. Also, we found that carrier frequency difference and location difference interacted with each other. This result contradicts the finding of independence between these two cues by Denham et al. (2010). It is likely that these interactions resulted from a ceiling effect: there was not enough room for further facilitating stream segregation by adding another cue.
Several studies indicated that rhythm perception doesn’t require musical training and the ability is present even in infancy (Demany et al., 1977; Honing et al., 2009; Winkler et al., 2009b). However, only few studies exist which investigated the effect rhythmic structure on stream segregation (but see Andreou et al., 2011; French-St George & Bregman, 1989;
Rimmele et al., 2012; Rogers & Bregman, 1993). Also, the effect of melody is not well understood. It was shown by Dowling and colleagues (Dowling, 1973; Dowling et al., 1987) that when a piece of familiar melody was presented to listeners and afterward this melody was presented together with distracting sounds from the same pitch range, participants were able to separate the different sound streams and recognize the familiar melody. These studies suggest that sequential stream segregation can depend on previous knowledge. Study II demonstrated that the presence of a melody facilitated switching towards segregation reducing the overall phase duration of the integrated percept and therefore utilized as a percept-inducing cue. However the effect of rhythm was due to the overlap between the two sets of sounds rather than the temporal arrangement of sounds within one set. An important finding is that the melodic pattern used in Study II induced stream segregation regardless of the familiarity. This result is contradictory to the results of Bendixen et al. (Bendixen et al., 2013;
Bendixen et al., 2010) who suggested that predictability-based cues such as temporal regularities can stabilize the sound streams but they do not induce stream segregation. In our case unfamiliar sound patterns were unpredictable as participants could not have predicted
99 upcoming sounds from the preceding ones. Similarly to our result Devergie et al. (Devergie et al., 2010) found that familiar melodies helped perceiving two stream without spectral differences between the two interleaved sequences. However in our experiment the effect was present even without familiarity of the melodies. Possible explanations of this result are discussed in detail in Study II. However, here we only wish to emphasize that Study II brought evidence against the notion that higher-order cues can only have percept stabilizing effects: we have shown that these cues can also induce stream segregation.
Percept-dependent processing of regular sounds
In Study IV we found that the amplitude of the P1 ERP component correlated with whether the sound was a part of the fore- or the background. When a sound was part of the background, it elicited a higher-amplitude P1 than when the same sound belonged to the foreground. In Study III the P1 was also enhanced when the percept was segregated for the first A and the B tones of the ABA triplets, whereas N1 was enhanced for the integrated percept. These results suggest that the perceived sound organization influences sound processing as early as 70 ms from stimulus onset.
Two different hypotheses have been put forward about the function of the P1 component. According to the baseline hypothesis, originally suggested by Luck et al. (Luck &
Hillyard, 1995; Luck et al., 1994) and Hillyard et al. (1998) the amplitude of the P1 decreases in unattended conditions compared to a neutral baseline. In contrast, the inhibition hypothesis proposes that the amplitude of the P1 increases with inhibition (Klimesch, 2011). This hypothesis suggests that the P1 doesn’t reflect sensory processing and cannot be explained as a sensory evoked component as it is not affected by the physical stimulus properties per se: P1 is only modulated by the stimulus properties when these features are relevant for early categorization or when the sounds are targets in the listeners’ task. Rather, P1 reflects
100 inhibition of task-irrelevant stimuli and/or networks. (Note, however that this hypothesis was put forward for the visual P1.) According to the inhibition hypothesis, larger P1 would be expected for the background sounds and not for the foreground sounds as the baseline hypothesis suggests. The results of Study IV are compatible with the inhibition hypothesis.
When participants perceived timbre A in the foreground, sounds with the timbre B acted as distractors (and vice versa). Therefore, perceiving a coherent falling A stream, B sounds might have been suppressed and, as a consequence, they elicited higher P1 compared to the falling A sounds. In contrast, in the rising percept we don’t find this difference between timbre A and B sounds as they constructed together the integrated percept (and no inhibition of the sounds was needed). However this interpretation is put into doubt by the results of Study III where the first A tone and the B tone of the ABA triplet elicited larger P1 when the percept was segregated. One possible explanation can be based on the fact that during the segregated percept, one of the two tones is part of the foreground, whereas the other is part of the background, and the foreground-background can switch both within and between segregated phases. If both the A and the B stream was perceived as background for sufficient proportion of time during the segregated phases (either by switching foreground and background within listeners or by some listeners predominantly perceiving one while others the other as background), then P1’s for both of these tones could have been enhanced at the group level.
Unfortunately, Study III provided no information regarding the selection of foreground within the segregated phases. It is also possible that the P1 modulations observed in the two studies may reflect partly different or interacting underlying effects, as the paradigms are not directly comparable. P1 is generally regarded as a component involved in feature extraction (see e.g., Näätänen & Winkler, 1999). Thus it is possible that the P1 enhancement reflects a modulation of feature extraction stream during segregation, or even the inhibition of this process for background sounds within the segregated sound organization. Therefore, further studies are
101 required regarding the P1 effects found in these two studies. Regardless of the precise theoretical interpretation, these studies showed that the percept influenced the sound processing as early as 70 ms from stimulus onset.
Percept-dependent processing of irregular sounds
There are several studies indicating that deviant sound processing follows the dominant percept (Rahne et al., 2007; Sussman et al., 1998, 1999; Winkler et al., 2003a;
Winkler et al., 2003b; Winkler et al., 2003c; Winkler et al., 2005). Most of these studies investigated the MMN elicitation corresponding to conscious perception (but see Winkler et al., 2005). However in the study of Ross et al. (1996) MMN elicitation did not match the listener’s percept. Rather, it followed the acoustic make-up of the sounds. Similarly, our results in Study IV showed that MMN elicitation did not follow the perceptual organization of the sounds. Rather, it followed the physical arrangement supporting that MMN is not directly related to auditory stream segregation. This result is consistent with results of other studies demonstrating dissociation between perception and MMN elicitation (Sussman et al., 2002;
Takegata et al., 2005; van Zuijen et al., 2006).
Some other ERP components displayed percept-related differences in processing irregular sounds. In Winkler et al.’ (2005) study, the auditory streaming paradigm was used with occasional sound omissions. They found an enhanced frontocentrally negative difference for the omission deviants when integration was perceived compared to the segregated percept.
The observed ERP component peaked at 176 ms from the onset of the expected tone. In Study III, we found similar effects for our frequency-deviant tones: Deviant tones encountered during experiencing the integrated percept elicited larger N2 than those encountered during the segregated percept. In contrast, the P3a component was larger during segregation than integration. This result might reflect the perceptual salience of the deviant tones in the two
102 cases. In a heterogeneous set of sounds (integrated streams contained two types of tones), a deviant sound may catch less attention, because a wider frequency range is covered by the auditory system. Both N2 and P3a are regarded as electrophysiological correlates of processes evaluating the contextual relevance of incoming sounds. Whereas the N2 component is regarded as an index of stimulus classification (Ritter et al., 1979), the P3a has been suggested to reflect attentional switching (Escera et al., 2000; Friedman et al., 2001; Polich, 2007;
Schröger, 1996), awareness of the participants that an unexpected event has occurred (Leppert et al., 2003), or contextual evaluation of novelty (Horváth et al., 2008). Our N2 finding might be explained by considering the position of the deviant tones. In Study III, deviants were always placed in the third position of the ABA triplets. Within this paradigm, when the sequence is perceived as integrated, the third tone defines the end of the repeating pattern. In contrast, when segregation is perceived, a continuous flow of sounds is heard and the information about the position is lost. Therefore deviation at the end of the triplets might be more relevant than in an arbitrary position within a homogeneous sequence. This could have caused the more negative N2 response for the integrated than for the segregated deviants. In summary irregular sound processing is strongly influenced by the current percept as was reflected by the N2 and P3a components.
A new stimulus paradigm for investigating auditory multistability
Study IV presented listeners with a multistable auditory stimulus sequence based on Wessel’s (1979) stimulus configuration. In this paradigm sounds of two different timbres were alternated. Within this paradigm, integration is defined as the perception of a coherent pattern composed of alternating timbres, whereas segregation as the perception of patterns with homogeneous timbres. The behavioral results found in Study IV showed similarities to studies employing variants of the auditory streaming paradigm (Bendixen et al., 2013;
Bendixen et al., 2010; Bőhm et al., 2013; Denham et al., 2014; Denham et al., 2010, 2013;
103 Pressnitzer & Hupé, 2006; Roberts et al., 2002). Participants reported perceptual switching throughout the stimulus blocks. Using timbre difference between the sounds, participants were able to easily decide which sound organization they perceived in the foreground.
Participants’ subjective report was that when they heard two streams only one of the timbres was perceived in the foreground. Therefore we suggest that this paradigm is a possible tool for investigating foreground-background discrimination with the advantages of bi-/multistable auditory stimulus configurations, that is, different perceptual foreground-background configurations without corresponding changes in the stimuli.
Conclusions
In conclusion, we found that both spectral and temporal-structure-based cues can induce auditory stream segregation. The fact that unfamiliar melodies induced stream segregation brings up the possibility that tonality analysis may influence auditory stream segregation. The nature of these processes requires further experiments. Our results showed multiple interactions between the processing of incoming sounds and the currently dominant representation of the sound sequence. Early effects of perceptual modulation in the ERP components reflected in the P1 and N1 occurred for regular sounds. Our results for the P1 suggest that enhancement of this component reflects inhibition of background tones rather than attention towards foreground tones. The processing of irregularity was affected in a later, evaluation stage, as reflected by modulations of the N2 and P3a ERP components. We have also tested a new auditory multistability paradigm for investigating the foreground-background decomposition of auditory scenes. With this paradigm, percept-dependent modulation of the P1 was obtained, yet MMN elicitation was independent of the percept.
Future studies should be conducted to clarify the relationship between the representations underlying the MMN ERP component and those involved in conscious perception.
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