Selective modulation of interhemispheric connectivity by transcranial alternating current stimulation influences binaural integration.

Brain connectivity plays a major role in the encoding, transfer, and integration of sensory information. Interregional synchronization of neural oscillations in the γ-frequency band has been suggested as a key mechanism underlying perceptual integration. In a recent study, we found evidence for this hypothesis showing that the modulation of interhemispheric oscillatory synchrony by means of bihemispheric high-density transcranial alternating current stimulation (HD-TACS) affects binaural integration of dichotic acoustic features. Here, we aimed to establish a direct link between oscillatory synchrony, effective brain connectivity, and binaural integration. We experimentally manipulated oscillatory synchrony (using bihemispheric γ-TACS with different interhemispheric phase lags) and assessed the effect on effective brain connectivity and binaural integration (as measured with functional MRI and a dichotic listening task, respectively). We found that TACS reduced intrahemispheric connectivity within the auditory cortices and antiphase (interhemispheric phase lag 180°) TACS modulated connectivity between the two auditory cortices. Importantly, the changes in intra- and interhemispheric connectivity induced by TACS were correlated with changes in perceptual integration. Our results indicate that γ-band synchronization between the two auditory cortices plays a functional role in binaural integration, supporting the proposed role of interregional oscillatory synchrony in perceptual integration.

Bilateral Gamma/Delta Transcranial Alternating Current Stimulation Affects Interhemispheric Speech Sound Integration.

Perceiving speech requires the integration of different speech cues, that is, formants. When the speech signal is split so that different cues are presented to the right and left ear (dichotic listening), comprehension requires the integration of binaural information. Based on prior electrophysiological evidence, we hypothesized that the integration of dichotically presented speech cues is enabled by interhemispheric phase synchronization between primary and secondary auditory cortex in the gamma frequency band. We tested this hypothesis by applying transcranial alternating current stimulation (TACS) bilaterally above the superior temporal lobe to induce or disrupt interhemispheric gamma-phase coupling. In contrast to initial predictions, we found that gamma TACS applied in-phase above the two hemispheres (interhemispheric lag 0°) perturbs interhemispheric integration of speech cues, possibly because the applied stimulation perturbs an inherent phase lag between the left and right auditory cortex. We also observed this disruptive effect when applying antiphasic delta TACS (interhemispheric lag 180°). We conclude that interhemispheric phase coupling plays a functional role in interhemispheric speech integration. The direction of this effect may depend on the stimulation frequency.

Hemispheric specializations affect interhemispheric speech sound integration during duplex perception.

The present study investigated whether speech-related spectral information benefits from initially predominant right or left hemisphere processing. Normal hearing individuals categorized speech sounds composed of an ambiguous base (perceptually intermediate between /ga/ and /da/), presented to one ear, and a disambiguating low or high F3 chirp presented to the other ear. Shorter response times were found when the chirp was presented to the left ear than to the right ear (inducing initially right-hemisphere chirp processing), but no between-ear differences in strength of overall integration. The results are in line with the assumptions of a right hemispheric dominance for spectral processing.

The interrelation between acoustic context effects and available response categories in speech sound categorization.

In an investigation of contextual influences on sound categorization, 64 Peruvian Spanish listeners categorized vowels on an /i/ to /e/ continuum. First, to measure the influence of the stimulus range (broad acoustic context) and the preceding stimuli (local acoustic context), listeners were presented with different subsets of the Spanish /i/-/e/ continuum in separate blocks. Second, the influence of the number of response categories was measured by presenting half of the participants with /i/ and /e/ as responses, and the other half with /i/, /e/, /a/, /o/, and /u/. The results showed that the perceptual category boundary between /i/ and /e/ shifted depending on the stimulus range and that the formant values of locally preceding items had a contrastive influence. Categorization was less susceptible to broad and local acoustic context effects, however, when listeners were presented with five rather than two response options. Vowel categorization depends not only on the acoustic properties of the target stimulus, but also on its broad and local acoustic context. The influence of such context is in turn affected by the number of internal referents that are available to the listener in a task.

Integral perception, but separate processing: The perceptual normalization of lexical tones and vowels.

In tonal languages, speech variability arises in both lexical tone (i.e., suprasegmentally) and vowel quality (segmentally). Listeners can use surrounding speech context to overcome variability in both speech cues, a process known as extrinsic normalization. Although vowels are the main carriers of tones, it is still unknown whether the combined percept (lexical tone and vowel quality) is normalized integrally or in partly separate processes. Here we used electroencephalography (EEG) to investigate the time course of lexical tone normalization and vowel normalization to answer this question. Cantonese adults listened to synthesized three-syllable stimuli in which the identity of a target syllable - ambiguous between high vs. mid-tone (Tone condition) or between /o/ vs. /u/ (Vowel condition) - was dependent on either the tone range (Tone condition) or the formant range (Vowel condition) of the first two syllables. It was observed that the ambiguous tone was more often interpreted as a high-level tone when the context had a relatively low pitch than when it had a high pitch (Tone condition). Similarly, the ambiguous vowel was more often interpreted as /o/ when the context had a relatively low formant range than when it had a relatively high formant range (Vowel condition). These findings show the typical pattern of extrinsic tone and vowel normalization. Importantly, the EEG results of participants showing the contrastive normalization effect demonstrated that the effects of vowel normalization could already be observed within the N2 time window (190-350 ms), while the first reliable effect of lexical tone normalization on cortical processing was observable only from the P3 time window (220-500 ms) onwards. The ERP patterns demonstrate that the contrastive perceptual normalization of lexical tones and that of vowels occur at least in partially separate time windows. This suggests that the extrinsic normalization can operate at the level of phonemes and tonemes separately instead of operating on the whole syllable at once.

Temporal contrast effects in human speech perception are immune to selective attention.

Two fundamental properties of perception are selective attention and perceptual contrast, but how these two processes interact remains unknown. Does an attended stimulus history exert a larger contrastive influence on the perception of a following target than unattended stimuli? Dutch listeners categorized target sounds with a reduced prefix "ge-" marking tense (e.g., ambiguous between gegaan-gaan "gone-go"). In 'single talker' Experiments 1-2, participants perceived the reduced syllable (reporting gegaan) when the target was heard after a fast sentence, but not after a slow sentence (reporting gaan). In 'selective attention' Experiments 3-5, participants listened to two simultaneous sentences from two different talkers, followed by the same target sounds, with instructions to attend only one of the two talkers. Critically, the speech rates of attended and unattended talkers were found to equally influence target perception - even when participants could watch the attended talker speak. In fact, participants' target perception in 'selective attention' Experiments 3-5 did not differ from participants who were explicitly instructed to divide their attention equally across the two talkers (Experiment 6). This suggests that contrast effects of speech rate are immune to selective attention, largely operating prior to attentional stream segregation in the auditory processing hierarchy.

Spectral contrast effects are modulated by selective attention in "cocktail party" settings.

Speech sounds are perceived relative to spectral properties of surrounding speech. For instance, target words that are ambiguous between /bɪt/ (with low F1) and /bɛt/ (with high F1) are more likely to be perceived as "bet" after a "low F1" sentence, but as "bit" after a "high F1" sentence. However, it is unclear how these spectral contrast effects (SCEs) operate in multi-talker listening conditions. Recently, Feng and Oxenham (J.Exp.Psychol.-Hum.Percept.Perform. 44(9), 1447-1457, 2018b) reported that selective attention affected SCEs to a small degree, using two simultaneously presented sentences produced by a single talker. The present study assessed the role of selective attention in more naturalistic "cocktail party" settings, with 200 lexically unique sentences, 20 target words, and different talkers. Results indicate that selective attention to one talker in one ear (while ignoring another talker in the other ear) modulates SCEs in such a way that only the spectral properties of the attended talker influences target perception. However, SCEs were much smaller in multi-talker settings (Experiment 2) than those in single-talker settings (Experiment 1). Therefore, the influence of SCEs on speech comprehension in more naturalistic settings (i.e., with competing talkers) may be smaller than estimated based on studies without competing talkers.

Initiation of utterance planning in response to pre-recorded and "live" utterances.

In everyday conversation, interlocutors often plan their utterances while listening to their conversational partners, thereby achieving short gaps between their turns. Important issues for current psycholinguistics are how interlocutors distribute their attention between listening and speech planning and how speech planning is timed relative to listening. Laboratory studies addressing these issues have used a variety of paradigms, some of which have involved using recorded speech to which participants responded, whereas others have involved interactions with confederates. This study investigated how this variation in the speech input affected the participants' timing of speech planning. In Experiment 1, participants responded to utterances produced by a confederate, who sat next to them and looked at the same screen. In Experiment 2, they responded to recorded utterances of the same confederate. Analyses of the participants' speech, their eye movements, and their performance in a concurrent tapping task showed that, compared with recorded speech, the presence of the confederate increased the processing load for the participants, but did not alter their global sentence planning strategy. These results have implications for the design of psycholinguistic experiments and theories of listening and speaking in dyadic settings.

Transformation of a temporal speech cue to a spatial neural code in human auditory cortex.

In speech, listeners extract continuously-varying spectrotemporal cues from the acoustic signal to perceive discrete phonetic categories. Spectral cues are spatially encoded in the amplitude of responses in phonetically-tuned neural populations in auditory cortex. It remains unknown whether similar neurophysiological mechanisms encode temporal cues like voice-onset time (VOT), which distinguishes sounds like /b/ and/p/. We used direct brain recordings in humans to investigate the neural encoding of temporal speech cues with a VOT continuum from /ba/ to /pa/. We found that distinct neural populations respond preferentially to VOTs from one phonetic category, and are also sensitive to sub-phonetic VOT differences within a population's preferred category. In a simple neural network model, simulated populations tuned to detect either temporal gaps or coincidences between spectral cues captured encoding patterns observed in real neural data. These results demonstrate that a spatial/amplitude neural code underlies the cortical representation of both spectral and temporal speech cues.

Speaker-normalized sound representations in the human auditory cortex.

The acoustic dimensions that distinguish speech sounds (like the vowel differences in "boot" and "boat") also differentiate speakers' voices. Therefore, listeners must normalize across speakers without losing linguistic information. Past behavioral work suggests an important role for auditory contrast enhancement in normalization: preceding context affects listeners' perception of subsequent speech sounds. Here, using intracranial electrocorticography in humans, we investigate whether and how such context effects arise in auditory cortex. Participants identified speech sounds that were preceded by phrases from two different speakers whose voices differed along the same acoustic dimension as target words (the lowest resonance of the vocal tract). In every participant, target vowels evoke a speaker-dependent neural response that is consistent with the listener's perception, and which follows from a contrast enhancement model. Auditory cortex processing thus displays a critical feature of normalization, allowing listeners to extract meaningful content from the voices of diverse speakers.

Lexical tone is perceived relative to locally surrounding context, vowel quality to preceding context.

Important speech cues such as lexical tone and vowel quality are perceptually contrasted to the distribution of those same cues in surrounding contexts. However, it is unclear whether preceding and following contexts have similar influences, and to what extent those influences are modulated by the auditory history of previous trials. To investigate this, Cantonese participants labeled sounds from (a) a tone continuum (mid- to high-level), presented with a context that had raised or lowered fundamental frequency (F0) values and (b) a vowel quality continuum (/u/ to /o/), where the context had raised or lowered first formant (F1) values. Contexts with high or low F0/F1 were presented in separate blocks or intermixed in 1 block. Contexts were presented following (Experiment 1) or preceding the target continuum (Experiment 2). Contrastive effects were found for both tone and vowel quality (e.g., decreased F0 values in contexts lead to more high tone target judgments and vice versa). Importantly, however, lexical tone was only influenced by F0 in immediately preceding and following contexts. Vowel quality was only influenced by the F1 in preceding contexts, but this extended to contexts from preceding trials. Contextual influences on tone and vowel quality are qualitatively different, which has important implications for understanding the mechanism of context effects in speech perception. (PsycINFO Database Record

Perceptual restoration of masked speech in human cortex.

Humans are adept at understanding speech despite the fact that our natural listening environment is often filled with interference. An example of this capacity is phoneme restoration, in which part of a word is completely replaced by noise, yet listeners report hearing the whole word. The neurological basis for this unconscious fill-in phenomenon is unknown, despite being a fundamental characteristic of human hearing. Here, using direct cortical recordings in humans, we demonstrate that missing speech is restored at the acoustic-phonetic level in bilateral auditory cortex, in real-time. This restoration is preceded by specific neural activity patterns in a separate language area, left frontal cortex, which predicts the word that participants later report hearing. These results demonstrate that during speech perception, missing acoustic content is synthesized online from the integration of incoming sensory cues and the internal neural dynamics that bias word-level expectation and prediction.

Variation in dual-task performance reveals late initiation of speech planning in turn-taking.

The smooth transitions between turns in natural conversation suggest that speakers often begin to plan their utterances while listening to their interlocutor. The presented study investigates whether this is indeed the case and, if so, when utterance planning begins. Two hypotheses were contrasted: that speakers begin to plan their turn as soon as possible (in our experiments less than a second after the onset of the interlocutor's turn), or that they do so close to the end of the interlocutor's turn. Turn-taking was combined with a finger tapping task to measure variations in cognitive load. We assumed that the onset of speech planning in addition to listening would be accompanied by deterioration in tapping performance. Two picture description experiments were conducted. In both experiments there were three conditions: (1) Tapping and Speaking, where participants tapped a complex pattern while taking over turns from a pre-recorded speaker, (2) Tapping and Listening, where participants carried out the tapping task while overhearing two pre-recorded speakers, and (3) Speaking Only, where participants took over turns as in the Tapping and Speaking condition but without tapping. The experiments differed in the amount of tapping training the participants received at the beginning of the session. In Experiment 2, the participants' eye-movements were recorded in addition to their speech and tapping. Analyses of the participants' tapping performance and eye movements showed that they initiated the cognitively demanding aspects of speech planning only shortly before the end of the turn of the preceding speaker. We argue that this is a smart planning strategy, which may be the speakers' default in many everyday situations.

Divide and conquer: How perceptual contrast sensitivity and perceptual learning cooperate in reducing input variation in speech perception.

Listeners have to overcome variability of the speech signal that can arise, for example, because of differences in room acoustics, differences in speakers' vocal tract properties, or idiosyncrasies in pronunciation. Two mechanisms that are involved in resolving such variation are perceptually contrastive effects that arise from surrounding acoustic context and lexically guided perceptual learning. Although both processes have been studied in great detail, little attention has been paid to how they operate relative to each other in speech perception. The present study set out to address this issue. The carrier parts of exposure stimuli of a classical perceptual learning experiment were spectrally filtered such that the acoustically ambiguous final fricatives sounded relatively more like the lexically intended sound (Experiment 1) or the alternative (Experiment 2). Perceptual learning was found only in the latter case. The findings show that perceptual contrast effects precede lexically guided perceptual learning, at least in terms of temporal order, and potentially in terms of cognitive processing levels as well.

Evidence for precategorical extrinsic vowel normalization.

Three experiments investigated whether extrinsic vowel normalization takes place largely at a categorical or a precategorical level of processing. Traditional vowel normalization effects in categorization were replicated in Experiment 1: Vowels taken from an [I]-[ε] continuum were more often interpreted as /I/ (which has a low first formant, F(1)) when the vowels were heard in contexts that had a raised F(1) than when the contexts had a lowered F(1). This was established with contexts that consisted of only two syllables. These short contexts were necessary for Experiment 2, a discrimination task that encouraged listeners to focus on the perceptual properties of vowels at a precategorical level. Vowel normalization was again found: Ambiguous vowels were more easily discriminated from an endpoint [ε] than from an endpoint [I] in a high-F(1) context, whereas the opposite was true in a low-F(1) context. Experiment 3 measured discriminability between pairs of steps along the [I]-[ε] continuum. Contextual influences were again found, but without discrimination peaks, contrary to what was predicted from the same participants' categorization behavior. Extrinsic vowel normalization therefore appears to be a process that takes place at least in part at a precategorical processing level.

Hemispheric differences in the effects of context on vowel perception.

Listeners perceive speech sounds relative to context. Contextual influences might differ over hemispheres if different types of auditory processing are lateralized. Hemispheric differences in contextual influences on vowel perception were investigated by presenting speech targets and both speech and non-speech contexts to listeners' right or left ears (contexts and targets either to the same or to opposite ears). Listeners performed a discrimination task. Vowel perception was influenced by acoustic properties of the context signals. The strength of this influence depended on laterality of target presentation, and on the speech/non-speech status of the context signal. We conclude that contrastive contextual influences on vowel perception are stronger when targets are processed predominately by the right hemisphere. In the left hemisphere, contrastive effects are smaller and largely restricted to speech contexts.

Constraints on the processes responsible for the extrinsic normalization of vowels.

Listeners tune in to talkers' vowels through extrinsic normalization. We asked here whether this process could be based on compensation for the long-term average spectrum (LTAS) of preceding sounds and whether the mechanisms responsible for normalization are indifferent to the nature of those sounds. If so, normalization should apply to nonspeech stimuli. Previous findings were replicated with first-formant (F1) manipulations of speech. Targets on a [pt]-[pɛt] (low-high F1) continuum were labeled as [pt] more after high-F1 than after low-F1 precursors. Spectrally rotated nonspeech versions of these materials produced similar normalization. None occurred, however, with nonspeech stimuli that were less speechlike, even though precursor-target LTAS relations were equivalent to those used earlier. Additional experiments investigated the roles of pitch movement, amplitude variation, formant location, and the stimuli's perceived similarity to speech. It appears that normalization is not restricted to speech but that the nature of the preceding sounds does matter. Extrinsic normalization of vowels is due, at least in part, to an auditory process that may require familiarity with the spectrotemporal characteristics of speech.

Listening to different speakers: on the time-course of perceptual compensation for vocal-tract characteristics.

This study used an active multiple-deviant oddball design to investigate the time-course of normalization processes that help listeners deal with between-speaker variability. Electroencephalograms were recorded while Dutch listeners heard sequences of non-words (standards and occasional deviants). Deviants were [ipapu] or [ɛpapu], and the standard was [(I)(ɛ)papu], where [(I)(ɛ)] was a vowel that was ambiguous between [ɛ] and [i]. These sequences were presented in two conditions, which differed with respect to the vocal-tract characteristics (i.e., the average 1st formant frequency) of the [papu] part, but not of the initial vowels [i], [ɛ] or [(I)(ɛ)] (these vowels were thus identical across conditions). Listeners more often detected a shift from [(I)(ɛ)papu] to [ɛpapu] than from [(I)(ɛ)papu] to [ipapu] in the high F(1) context condition; the reverse was true in the low F(1) context condition. This shows that listeners' perception of vowels differs depending on the speaker's vocal-tract characteristics, as revealed in the speech surrounding those vowels. Cortical electrophysiological responses reflected this normalization process as early as about 120 ms after vowel onset, which suggests that shifts in perception precede influences due to conscious biases or decision strategies. Listeners' abilities to normalize for speaker-vocal-tract properties are for an important part the result of a process that influences representations of speech sounds early in the speech processing stream.

The bounds on flexibility in speech perception.

Dutch listeners were exposed to the English theta sound (as in bath), which replaced [f] in /f/-final Dutch words or, for another group, [s] in /s/-final words. A subsequent identity-priming task showed that participants had learned to interpret theta as, respectively, /f/ or /s/. Priming effects were equally strong when the exposure sound was an ambiguous [fs]-mixture and when primes contained unambiguous fricatives. When the exposure sound was signal-correlated noise, listeners interpreted it as the spectrally similar /f/, irrespective of lexical bias during exposure. Perceptual learning about speech is thus constrained by spectral similarity between the input and established phonological categories, but within those limits, adjustments are thorough enough that even nonnative sounds can be treated fully as native sounds.