Exposure to bilingual or monolingual maternal speech during pregnancy affects the neurophysiological encoding of speech sounds in neonates differently.

Introduction: Exposure to maternal speech during the prenatal period shapes speech perception and linguistic preferences, allowing neonates to recognize stories heard frequently in utero and demonstrating an enhanced preference for their mother's voice and native language. Yet, with a high prevalence of bilingualism worldwide, it remains an open question whether monolingual or bilingual maternal speech during pregnancy influence differently the fetus' neural mechanisms underlying speech sound encoding. Methods: In the present study, the frequency-following response (FFR), an auditory evoked potential that reflects the complex spectrotemporal dynamics of speech sounds, was recorded to a two-vowel /oa/ stimulus in a sample of 129 healthy term neonates within 1 to 3 days after birth. Newborns were divided into two groups according to maternal language usage during the last trimester of gestation (monolingual; bilingual). Spectral amplitudes and spectral signal-to-noise ratios (SNR) at the stimulus fundamental (F0) and first formant (F1) frequencies of each vowel were, respectively, taken as measures of pitch and formant structure neural encoding. Results: Our results reveal that while spectral amplitudes at F0 did not differ between groups, neonates from bilingual mothers exhibited a lower spectral SNR. Additionally, monolingually exposed neonates exhibited a higher spectral amplitude and SNR at F1 frequencies. Discussion: We interpret our results under the consideration that bilingual maternal speech, as compared to monolingual, is characterized by a greater complexity in the speech sound signal, rendering newborns from bilingual mothers more sensitive to a wider range of speech frequencies without generating a particularly strong response at any of them. Our results contribute to an expanding body of research indicating the influence of prenatal experiences on language acquisition and underscore the necessity of including prenatal language exposure in developmental studies on language acquisition, a variable often overlooked yet capable of influencing research outcomes.

Neonatal Frequency-Following Responses: A Methodological Framework for Clinical Applications.

The frequency-following response (FFR) to periodic complex sounds is a noninvasive scalp-recorded auditory evoked potential that reflects synchronous phase-locked neural activity to the spectrotemporal components of the acoustic signal along the ascending auditory hierarchy. The FFR has gained recent interest in the fields of audiology and auditory cognitive neuroscience, as it has great potential to answer both basic and applied questions about processes involved in sound encoding, language development, and communication. Specifically, it has become a promising tool in neonates, as its study may allow both early identification of future language disorders and the opportunity to leverage brain plasticity during the first 2 years of life, as well as enable early interventions to prevent and/or ameliorate sound and language encoding disorders. Throughout the present review, we summarize the state of the art of the neonatal FFR and, based on our own extensive experience, present methodological approaches to record it in a clinical environment. Overall, the present review is the first one that comprehensively focuses on the neonatal FFRs applications, thus supporting the feasibility to record the FFR during the first days of life and the predictive potential of the neonatal FFR on detecting short- and long-term language abilities and disruptions.

Neural generators of the frequency-following response elicited to stimuli of low and high frequency: A magnetoencephalographic (MEG) study.

The frequency-following response (FFR) to periodic complex sounds has gained recent interest in auditory cognitive neuroscience as it captures with great fidelity the tracking accuracy of the periodic sound features in the ascending auditory system. Seminal studies suggested the FFR as a correlate of subcortical sound encoding, yet recent studies aiming to locate its sources challenged this assumption, demonstrating that FFR receives some contribution from the auditory cortex. Based on frequency-specific phase-locking capabilities along the auditory hierarchy, we hypothesized that FFRs to higher frequencies would receive less cortical contribution than those to lower frequencies, hence supporting a major subcortical involvement for these high frequency sounds. Here, we used a magnetoencephalographic (MEG) approach to trace the neural sources of the FFR elicited in healthy adults (N = 19) to low (89 Hz) and high (333 Hz) frequency sounds. FFRs elicited to the high and low frequency sounds were clearly observable on MEG and comparable to those obtained in simultaneous electroencephalographic recordings. Distributed source modeling analyses revealed midbrain, thalamic, and cortical contributions to FFR, arranged in frequency-specific configurations. Our results showed that the main contribution to the high-frequency sound FFR originated in the inferior colliculus and the medial geniculate body of the thalamus, with no significant cortical contribution. In contrast, the low-frequency sound FFR had a major contribution located in the auditory cortices, and also received contributions originating in the midbrain and thalamic structures. These findings support the multiple generator hypothesis of the FFR and are relevant for our understanding of the neural encoding of sounds along the auditory hierarchy, suggesting a hierarchical organization of periodicity encoding.

Emergence of prediction error along the human auditory hierarchy.

Auditory prediction errors have been extensively associated with the mismatch negativity (MMN), a cortical auditory evoked potential that denotes deviance detection. Yet, many studies lacked the appropriate controls to disentangle sensory adaptation from prediction error. Furthermore, subcortical deviance detection has been shown in humans through recordings of the frequency-following response (FFR), an early auditory evoked potential that reflects the neural tracking of the periodic characteristics of a sound, suggesting the possibility that prediction errors emerge subcortically in the auditory pathway. The present study aimed at investigating the emergence of prediction error along the auditory hierarchy in humans through combined recordings of the FFR and the MMN, tapping at subcortical and cortical levels, respectively, while disentangling prediction error from sensory adaptation with the use of appropriate controls. "Oddball" sequences of pure tones featuring repeated "standard" stimuli (269 Hz; p = 0.8) and rare "deviant" stimuli (p = 0.2; of 289, 329 and 409 Hz delivered in separated blocks to test "frequency separation" effects) were presented to nineteen healthy young participants. "Reversed" oddball sequences (where standard and deviant tones swapped their roles) were presented allowing comparison of responses to same physical stimuli as a function of functional role (i.e., standard, deviant). Critically, control sequences featuring five equiprobable tones (p = 0.2) allowed to dissociate sensory adaptation from prediction error. Results revealed that the MMN amplitude increased as a function of frequency separation yet displayed the same amplitude when retrieved against the control sequences, confirming previous results. FFRs showed repetition enhancement effects across all frequency separations, as supported by larger spectral amplitude to standard than to deviant and control stimuli. This pattern of results provides insights into the hierarchy of the human prediction error system in audition, suggesting that prediction errors in humans emerge at cortical level.

Effects of cTBS on the Frequency-Following Response and Other Auditory Evoked Potentials.

The frequency-following response (FFR) is an auditory evoked potential (AEP) that follows the periodic characteristics of a sound. Despite being a widely studied biosignal in auditory neuroscience, the neural underpinnings of the FFR are still unclear. Traditionally, FFR was associated with subcortical activity, but recent evidence suggested cortical contributions which may be dependent on the stimulus frequency. We combined electroencephalography (EEG) with an inhibitory transcranial magnetic stimulation protocol, the continuous theta burst stimulation (cTBS), to disentangle the cortical contribution to the FFR elicited to stimuli of high and low frequency. We recorded FFR to the syllable /ba/ at two fundamental frequencies (Low: 113 Hz; High: 317 Hz) in healthy participants. FFR, cortical potentials, and auditory brainstem response (ABR) were recorded before and after real and sham cTBS in the right primary auditory cortex. Results showed that cTBS did not produce a significant change in the FFR recorded, in any of the frequencies. No effect was observed in the ABR and cortical potentials, despite the latter known contributions from the auditory cortex. Possible reasons behind the negative results include compensatory mechanisms from the non-targeted areas, intraindividual variability of the cTBS effectiveness, and the particular location of our target area, the primary auditory cortex.

Phonological Task Enhances the Frequency-Following Response to Deviant Task-Irrelevant Speech Sounds.

In electroencephalography (EEG) measurements, processing of periodic sounds in the ascending auditory pathway generates the frequency-following response (FFR) phase-locked to the fundamental frequency (F0) and its harmonics of a sound. We measured FFRs to the steady-state (vowel) part of syllables /ba/ and /aw/ occurring in binaural rapid streams of speech sounds as frequently repeating standard syllables or as infrequent (p = 0.2) deviant syllables among standard /wa/ syllables. Our aim was to study whether concurrent active phonological processing affects early processing of irrelevant speech sounds reflected by FFRs to these sounds. To this end, during syllable delivery, our healthy adult participants performed tasks involving written letters delivered on a computer screen in a rapid stream. The stream consisted of vowel letters written in red, infrequently occurring consonant letters written in the same color, and infrequently occurring vowel letters written in blue. In the phonological task, the participants were instructed to press a response key to the consonant letters differing phonologically but not in color from the frequently occurring red vowels, whereas in the non-phonological task, they were instructed to respond to the vowel letters written in blue differing only in color from the frequently occurring red vowels. We observed that the phonological task enhanced responses to deviant /ba/ syllables but not responses to deviant /aw/ syllables. This suggests that active phonological task performance may enhance processing of such small changes in irrelevant speech sounds as the 30-ms difference in the initial formant-transition time between the otherwise identical syllables /ba/ and /wa/ used in the present study.

The Potential Effect of Forbrain as an Altered Auditory Feedback Device.

Purpose: The purpose of this study was to run a proof of concept on a new commercially available device, Forbrain® (Sound For Life Ltd/Soundev, Luxemburg, model UN38.3), to test whether it can modulate the speech of its users. Method: Participants were instructed to read aloud a text of their choice during 3 experimental phases: baseline, test, and posttest, while wearing a Forbrain® headset. Critically, for half of the participants (Forbrain group), the device was turned on during the test phase, whereas for the other half (control group), the device was kept off. Voice recordings were analyzed to derive 6 quantitative measures of voice quality over each of the phases of the experiment. Results: A significant Group × Phase interaction was obtained for the smoothed cepstral peak prominence, a measure of voice harmony, and for the trendline of the long-term average spectrum, a measure of voice robustness, this latter surviving Bonferroni correction for multiple comparisons. Conclusions: The results of this study indicate the effectiveness of Forbrain® in modifying the speech of its users. It is suggested that Forbrain® works as an altered auditory feedback device. It may hence be used as a clinical device in speech therapy clinics, yet further studies are warranted to test its usefulness in clinical groups.

Timing predictability enhances regularity encoding in the human subcortical auditory pathway.

The encoding of temporal regularities is a critical property of the auditory system, as short-term neural representations of environmental statistics serve to auditory object formation and detection of potentially relevant novel stimuli. A putative neural mechanism underlying regularity encoding is repetition suppression, the reduction of neural activity to repeated stimulation. Although repetitive stimulation per se has shown to reduce auditory neural activity in animal cortical and subcortical levels and in the human cerebral cortex, other factors such as timing may influence the encoding of statistical regularities. This study was set out to investigate whether temporal predictability in the ongoing auditory input modulates repetition suppression in subcortical stages of the auditory processing hierarchy. Human auditory frequency-following responses (FFR) were recorded to a repeating consonant-vowel stimuli (/wa/) delivered in temporally predictable and unpredictable conditions. FFR amplitude was attenuated by repetition independently of temporal predictability, yet we observed an accentuated suppression when the incoming stimulation was temporally predictable. These findings support the view that regularity encoding spans across the auditory hierarchy and point to temporal predictability as a modulatory factor of regularity encoding in early stages of the auditory pathway.