Makio Kashino

Involvement of the Thalamocortical Loop in the Spontaneous Switching of Percepts in Auditory Streaming

Perceptual grouping of successive frequency components, namely, auditory streaming, is essential for auditory scene analysis. Prolonged listening to an unchanging triplet-tone sequence produces a series of illusory switches between a single coherent stream (S1) and two distinct streams (S2). The predominant percept depends on the frequency difference (Δf) between high and low tones. Here, we combined the use of different Δfs with an event-related fMRI design to identify whether the temporal dynamics of brain activity differs depending on the direction of perceptual switches. The results demonstrated that the activity of the medial geniculate body (MGB) in the thalamus occurred earlier during switching from nonpredominant to predominant percepts, whereas that of the auditory cortex (AC) occurred earlier during switching from predominant to nonpredominant percepts, regardless of Δf. The asymmetry of temporal precedence indicates that the MGB and AC activations play different roles in perceptual switching and depend on perceptual predominance rather than on S1 and S2 percepts per se. Our results suggest that feedforward and feedback processes in the thalamocortical loop are involved in the formation of percepts in auditory streaming.

Structure of Rat Ultrasonic Vocalizations and Its Relevance to Behavior

Rats are known to emit ultrasonic vocalizations (USVs). These USVs have been hypothesized to hold biological meaning, and the relationship between USVs and behavior has been extensively studied. However, most of these studies looked at specific conditions, such as fear-inducing situations and sexual encounters. In the present experiment, the USVs of pairs of rats in ordinary housing conditions were recorded and their features were examined. Three clusters of USVs in the 25-, 40-, and 60-kHz range were detected, which roughly corresponded to fighting, feeding, and moving, respectively. We analyzed sequential combinations of two or more clusters using a state transition model. The results revealed a more specific correspondence between the USVs and behaviors, suggesting that rat USV may work as a type of communication tool.

The role of temporal regularity in auditory segregation.

The idea that predictive modelling and extraction of regularities plays a pivotal role in auditory segregation has recently attracted considerable attention. The present study investigated the effect of one basic form of regularity, rhythmic regularity, on auditory stream segregation. We departed from the classic streaming paradigm and developed a new stimulus, Rand-AB, consisting of two, concurrently presented, temporally uncorrelated, tone sequences (with frequencies A and B). To evaluate segregation, we used an objective measure of the extent to which listeners are able to selectively attend to one of the sequences in the presence of the other. Performance was quantified on a difficult pattern detection task which involves detecting a rarely occurring pattern of amplitude modulation applied to three consecutive A or B tones. In all cases the attended sequence was temporally irregular (with a random inter-tone-interval (ITI) between 100 and 400 ms) and the regularity status of the competing sequence was set to one of four conditions: (1) random ITI between 100 and 400 ms (2) isochronous with ITI = 400 ms. (3) isochronous with ITI = 250 ms (equal to the mean rate of the attended sequence) (4) isochronous with ITI = 100 ms. For a frequency separation of 2 (but not 4) semi tones we observed improved performance in conditions (3) and (4) relative to (1), suggesting that stream segregation is facilitated when the distracter sequence is temporally regular, but that the effect of temporal regularity as a cue for segregation is limited to relatively fast rates and to situations where frequency separation is insufficient for segregation. These findings provide new evidence to support models of streaming that involve segregation based on the formation of predictive models.

Functional brain networks underlying perceptual switching: auditory streaming and verbal transformations

Recent studies have shown that auditory scene analysis involves distributed neural sites below, in, and beyond the auditory cortex (AC). However, it remains unclear what role each site plays and how they interact in the formation and selection of auditory percepts. We addressed this issue through perceptual multistability phenomena, namely, spontaneous perceptual switching in auditory streaming (AS) for a sequence of repeated triplet tones, and perceptual changes for a repeated word, known as verbal transformations (VTs). An event-related fMRI analysis revealed brain activity timelocked to perceptual switching in the cerebellum for AS, in frontal areas for VT, and the AC and thalamus for both. The results suggest that motor-based prediction, produced by neural networks outside the auditory system, plays essential roles in the segmentation of acoustic sequences both in AS and VT. The frequency of perceptual switching was determined by a balance between the activation of two sites, which are proposed to be involved in exploring novel perceptual organization and stabilizing current perceptual organization. The effect of the gene polymorphism of catechol-O-methyltransferase (COMT) on individual variations in switching frequency suggests that the balance of exploration and stabilization is modulated by catecholamines such as dopamine and noradrenalin. These mechanisms would support the noteworthy flexibility of auditory scene analysis.

Neural mechanisms of auditory awareness underlying verbal transformations

Prolonged listening to a repeated word without a pause produces a series of illusory transitions of the physically unchanging word, which is called verbal transformation. Verbal transformations provide a rare opportunity to examine how auditory percepts are formed in the brain. We found that verbal forms are affected by phonetic reorganization of a word, rather than by auditory adaptation and lexical distortion of it. We identified brain activity leading to individual differences between perceptual transitions and tone detection. An event-related fMRI analysis revealed that the left inferior frontal cortex (IFC), anterior cingulate cortex (ACC), and the left prefrontal cortex were activated when perceptual transitions from one verbal form to another occurred, but not when tone pips were detected. The number of perceptual transitions showed positive and negative correlations with signal intensity in the left IFC and the left ACC, respectively. The results suggest that active generation of verbal forms is linked with articulatory gestures for speech production and that the frequency of perceptual transitions is determined by a balance of the activations between the two brain regions. Structural equation modeling demonstrated that individual differences in the number of perceptual transitions rely on negative feedback from the ACC to the IFC via the posterior insula. These findings suggest that distributed frontal areas are involved in auditory awareness underlying verbal transformations.

The role of spectral change detectors in temporal order judgment of tones.

Human listeners can judge the temporal order of acoustic events quite accurately in certain conditions. We hypothesized that this accuracy is realized by coding temporal order as a single neural code of spectral change in the auditory system. To test this hypothesis, we examined whether adaptation to linear frequency glides affects subsequent temporal order judgment of brief tones. We found that the point of subjective simultaneity between two tones shifted depending on the direction of spectral change of the adaptor, as predicted by the hypothesis. The amount of aftereffect was significantly reduced when adaptor and test tones were presented in different frequency regions or to different ears, suggesting that the relevant neural units exist predominantly in a frequency-selective monaural pathway.

One, two, many—Judging the number of concurrent talkers.

The ability of listeners to judge the number of concurrent talkers was examined. Ten female and 11 male Japanese talkers each recorded 20 familiar Japanese words consisting of four consonant–vowel syllables each. In each trial, a number of different talkers was chosen randomly from the same‐sex group, and presented synchronously to four native Japanese listeners, who were asked to judge how many talkers were present. The range of talker numbers was unknown to the listeners. To eliminate cues associated with level, the over‐all sound pressure level was varied randomly in each trial, with RMS levels of the individual words equalized. It was found that judgments were nearly perfect for up to two talkers, but deteriorated abruptly for three or more talkers. In the latter case, the number of talkers was underestimated, although estimates increased slightly as the number of talkers increased. Factors that may promote sound source separation, such as lexicality (e.g., forward versus reverse speech) and spatial s...

Effects of self-motion on auditory scene analysis

Auditory scene analysis requires the listener to parse the incoming flow of acoustic information into perceptual “streams,” such as sentences from a single talker in the midst of background noise. Behavioral and neural data show that the formation of streams is not instantaneous; rather, streaming builds up over time and can be reset by sudden changes in the acoustics of the scene. Here, we investigated the effect of changes induced by voluntary head motion on streaming. We used a telepresence robot in a virtual reality setup to disentangle all potential consequences of head motion: changes in acoustic cues at the ears, changes in apparent source location, and changes in motor or attentional processes. The results showed that self-motion influenced streaming in at least two ways. Right after the onset of movement, self-motion always induced some resetting of perceptual organization to one stream, even when the acoustic scene itself had not changed. Then, after the motion, the prevalent organization was rapidly biased by the binaural cues discovered through motion. Auditory scene analysis thus appears to be a dynamic process that is affected by the active sensing of the environment.

Mechanisms Underlying Referral of Thermal Sensations to Sites of Tactile Stimulation

When three stimulators are simultaneously touched with the middle three fingers of one hand but only the outer two stimulators are cooled or heated, the central (neutral) stimulator is also perceived to be cold or warm. This phenomenon is known as thermal referral and it shares phenomenological similarities with filling-in, in which the discontinuity in the signals of interest can be compensated perceptually on the basis of the spatially adjacent context. Although the mechanisms underlying filling-in have been well substantiated, those underlying thermal referral are still poorly understood. In the present study, we examined the intensity perception of the sensation resulting from thermal referral with human participants. We found that the sensation was uniform among the three fingers, but its apparent intensity was always lower than the physical intensity applied to the outer two fingers. These results indicate that the thermal uniformity perceived under thermal referral is not created by the brains interpolating the thermal changes applied to the outer two fingers, as one would expect for those induced by typical filling-in. Instead, the thermal changes applied to the outer two fingers are summated and redistributed to all the fingers in contact. Our findings suggest that thermal referral is mediated by two separate processes. One determines the apparent intensity from the physical intensity and the areal extent of the thermal stimulation; the other determines the localization of the resulting sensation from the apparent sites of tactile stimulation.

The Tactile Continuity Illusion.

We can perceive the continuity of an object or event by integrating spatially/temporally discrete sensory inputs. The mechanism underlying this perception of continuity has intrigued many researchers and has been well documented in both the visual and auditory modalities. The present study shows for the first time to our knowledge that an illusion of continuity also occurs with vibrotactile stimulation. We found that when the brief temporal gaps inserted into a vibrotactile target were filled with vibrotactile noise, the target vibration was perceived to continue through the noise if the target vibration was sufficiently weak relative to the noise. It is important that the illusory continuity of the vibration cannot be distinguished from the physically continuous vibration. These results therefore suggest that the continuity illusion is common to multiple sensory modalities and that it reflects a fundamental principle of perception.