Neural indices of listening effort in noisy environments.

Listening in a noisy environment is challenging for individuals with normal hearing and can be a significant burden for those with hearing impairment. The extent to which this burden is alleviated by a hearing device is a major, unresolved issue for rehabilitation. Here, we found adult users of cochlear implants (CIs) self-reported listening effort during a speech-in-noise task that was positively related to alpha oscillatory activity in the left inferior frontal cortex, canonical Broca's area, and inversely related to speech envelope coherence in the 2-5 Hz range originating in the superior-temporal plane encompassing auditory cortex. Left frontal cortex coherence in the 2-5 Hz range also predicted speech-in-noise identification. These data demonstrate that neural oscillations predict both speech perception ability in noise and listening effort.

Region-specific modulations in oscillatory alpha activity serve to facilitate processing in the visual and auditory modalities.

There have been a number of studies suggesting that oscillatory alpha activity (~10 Hz) plays a pivotal role in attention by gating information flow to relevant sensory regions. The vast majority of these studies have looked at shifts of attention in the spatial domain and only in a single modality (often visual or sensorimotor). In the current magnetoencephalography (MEG) study, we investigated the role of alpha activity in the suppression of a distracting modality stream. We used a cross-modal attention task where visual cues indicated whether participants had to judge a visual orientation or discriminate the auditory pitch of an upcoming target. The visual and auditory targets were presented either simultaneously or alone, allowing us to behaviorally gauge the "cost" of having a distractor present in each modality. We found that the preparation for visual discrimination (relative to pitch discrimination) resulted in a decrease of alpha power (9-11 Hz) in the early visual cortex, with a concomitant increase in alpha/beta power (14-16 Hz) in the supramarginal gyrus, a region suggested to play a vital role in short-term storage of pitch information (Gaab et al., 2003). On a trial-by-trial basis, alpha power over the visual areas was significantly correlated with increased visual discrimination times, whereas alpha power over the precuneus and right superior temporal gyrus was correlated with increased auditory discrimination times. However, these correlations were only significant when the targets were paired with distractors. Our work adds to increasing evidence that the top-down (i.e. attentional) modulation of alpha activity is a mechanism by which stimulus processing can be gated within the cortex. Here, we find that this phenomenon is not restricted to the domain of spatial attention and can be generalized to other sensory modalities than vision.

Loudness adaptation accompanying ribbon synapse and auditory nerve disorders.

Abnormal auditory adaptation is a standard clinical tool for diagnosing auditory nerve disorders due to acoustic neuromas. In the present study we investigated auditory adaptation in auditory neuropathy owing to disordered function of inner hair cell ribbon synapses (temperature-sensitive auditory neuropathy) or auditory nerve fibres. Subjects were tested when afebrile for (i) psychophysical loudness adaptation to comfortably-loud sustained tones; and (ii) physiological adaptation of auditory brainstem responses to clicks as a function of their position in brief 20-click stimulus trains (#1, 2, 3 … 20). Results were compared with normal hearing listeners and other forms of hearing impairment. Subjects with ribbon synapse disorder had abnormally increased magnitude of loudness adaptation to both low (250 Hz) and high (8000 Hz) frequency tones. Subjects with auditory nerve disorders had normal loudness adaptation to low frequency tones; all but one had abnormal adaptation to high frequency tones. Adaptation was both more rapid and of greater magnitude in ribbon synapse than in auditory nerve disorders. Auditory brainstem response measures of adaptation in ribbon synapse disorder showed Wave V to the first click in the train to be abnormal both in latency and amplitude, and these abnormalities increased in magnitude or Wave V was absent to subsequent clicks. In contrast, auditory brainstem responses in four of the five subjects with neural disorders were absent to every click in the train. The fifth subject had normal latency and abnormally reduced amplitude of Wave V to the first click and abnormal or absent responses to subsequent clicks. Thus, dysfunction of both synaptic transmission and auditory neural function can be associated with abnormal loudness adaptation and the magnitude of the adaptation is significantly greater with ribbon synapse than neural disorders.

Auditory cortical activity in normal hearing subjects to consonant vowels presented in quiet and in noise.

OBJECTIVE: Compare brain potentials to consonant vowels (CVs) as a function of both voice onset times (VOTs) and consonant position; initial (CV) versus second (VCV). METHODS: Auditory cortical potentials (N100, P200, N200, and a late slow negativity, (SN) were recorded from scalp electrodes in twelve normal hearing subjects to consonant vowels in initial position (CVs: /du/ and /tu/), in second position (VCVs: /udu/ and /utu/), and to vowels alone (V: /u/) and paired (VVs: /uu/) separated in time to simulate consonant voice onset times (VOTs). RESULTS: CVs evoked "acoustic onset" N100s of similar latency but larger amplitudes to /du/ than /tu/. CVs preceded by a vowel (VCVs) evoked "acoustic change" N100s with longer latencies to /utu/ than /udu/. Their absolute latency difference was less than the corresponding VOT difference. The SN following N100 to VCVs was larger to /utu/ than /udu/. Paired vowels (/uu/) separated by intervals corresponding to consonant VOTs evoked N100s with latency differences equal to the simulated VOT differences and SNs of similar amplitudes. Noise masking resulted in VCV N100 latency differences that were now equal to consonant VOT differences. Brain activations by CVs, VCVs, and VVs were maximal in right temporal lobe. CONCLUSION: Auditory cortical activities to CVs are sensitive to: (1) position of the CV in the utterance; (2) VOTs of consonants; and (3) noise masking. SIGNIFICANCE: VOTs of stop consonants affect auditory cortical activities differently as a function of the position of the consonant in the utterance.

Towards a closed-loop cochlear implant system: application of embedded monitoring of peripheral and central neural activity.

Although the cochlear implant (CI) is widely considered the most successful neural prosthesis, it is essentially an open-loop system that requires extensive initial fitting and frequent tuning to maintain a high, but not necessarily optimal, level of performance. Two developments in neuroscience and neuroengineering now make it feasible to design a closed-loop CI. One development is the recording and interpretation of evoked potentials (EPs) from the peripheral to the central nervous system. The other is the embedded hardware and software of a modern CI that allows recording of EPs. We review EPs that are pertinent to behavioral functions from simple signal detection and loudness growth to speech discrimination and recognition. We also describe signal processing algorithms used for electric artifact reduction and cancellation, critical to the recording of electric EPs. We then present a conceptual design for a closed-loop CI that utilizes in an innovative way the embedded implant receiver and stimulators to record short latency compound action potentials ( ~1 ms), auditory brainstem responses (1-10 ms) and mid-to-late cortical potentials (20-300 ms). We compare EPs recorded using the CI to EPs obtained using standard scalp electrodes recording techniques. Future applications and capabilities are discussed in terms of the development of a new generation of closed-loop CIs and other neural prostheses.

Tinnitus suppression by low-rate electric stimulation and its electrophysiological mechanisms.

Tinnitus is a phantom sensation of sound in the absence of external stimulation. However, external stimulation, particularly electric stimulation via a cochlear implant, has been shown to suppress tinnitus. Different from traditional methods of delivering speech sounds or high-rate (>2000 Hz) stimulation, the present study found a unique unilaterally-deafened cochlear implant subject whose tinnitus was completely suppressed by a low-rate (<100 Hz) stimulus, delivered at a level softer than tinnitus to the apical part of the cochlea. Taking advantage of this novel finding, the present study compared both event-related and spontaneous cortical activities in the same subject between the tinnitus-present and tinnitus-suppressed states. Compared with the results obtained in the tinnitus-present state, the low-rate stimulus reduced cortical N100 potentials while increasing the spontaneous alpha power in the auditory cortex. These results are consistent with previous neurophysiological studies employing subjects with and without tinnitus and shed light on both tinnitus mechanism and treatment.

Auditory cortical N100 in pre- and post-synaptic auditory neuropathy to frequency or intensity changes of continuous tones.

OBJECTIVES: Auditory cortical N100s were examined in ten auditory neuropathy (AN) subjects as objective measures of impaired hearing. METHODS: Latencies and amplitudes of N100 in AN to increases of frequency (4-50%) or intensity (4-8 dB) of low (250 Hz) or high (4000 Hz) frequency tones were compared with results from normal-hearing controls. The sites of auditory nerve dysfunction were pre-synaptic (n=3) due to otoferlin mutations causing temperature sensitive deafness, post-synaptic (n=4) affecting other cranial and/or peripheral neuropathies, and undefined (n=3). RESULTS: AN consistently had N100s only to the largest changes of frequency or intensity whereas controls consistently had N100s to all but the smallest frequency and intensity changes. N100 latency in AN was significantly delayed compared to controls, more so for 250 than for 4000 Hz and more so for changes of intensity compared to frequency. N100 amplitudes to frequency change were significantly reduced in ANs compared to controls, except for pre-synaptic AN in whom amplitudes were greater than controls. N100 latency to frequency change of 250 but not of 4000 Hz was significantly related to speech perception scores. CONCLUSIONS: As a group, AN subjects' N100 potentials were abnormally delayed and smaller, particularly for low frequency. The extent of these abnormalities differed between pre- and post-synaptic forms of the disorder. SIGNIFICANCE: Abnormalities of auditory cortical N100 in AN reflect disorders of both temporal processing (low frequency) and neural adaptation (high frequency). Auditory N100 latency to the low frequency provides an objective measure of the degree of impaired speech perception in AN.

Cortical evoked potentials to an auditory illusion: binaural beats.

OBJECTIVE: To define brain activity corresponding to an auditory illusion of 3 and 6Hz binaural beats in 250Hz or 1000Hz base frequencies, and compare it to the sound onset response. METHODS: Event-Related Potentials (ERPs) were recorded in response to unmodulated tones of 250 or 1000Hz to one ear and 3 or 6Hz higher to the other, creating an illusion of amplitude modulations (beats) of 3Hz and 6Hz, in base frequencies of 250Hz and 1000Hz. Tones were 2000ms in duration and presented with approximately 1s intervals. Latency, amplitude and source current density estimates of ERP components to tone onset and subsequent beats-evoked oscillations were determined and compared across beat frequencies with both base frequencies. RESULTS: All stimuli evoked tone-onset P(50), N(100) and P(200) components followed by oscillations corresponding to the beat frequency, and a subsequent tone-offset complex. Beats-evoked oscillations were higher in amplitude with the low base frequency and to the low beat frequency. Sources of the beats-evoked oscillations across all stimulus conditions located mostly to left lateral and inferior temporal lobe areas in all stimulus conditions. Onset-evoked components were not different across stimulus conditions; P(50) had significantly different sources than the beats-evoked oscillations; and N(100) and P(200) sources located to the same temporal lobe regions as beats-evoked oscillations, but were bilateral and also included frontal and parietal contributions. CONCLUSIONS: Neural activity with slightly different volley frequencies from left and right ear converges and interacts in the central auditory brainstem pathways to generate beats of neural activity to modulate activities in the left temporal lobe, giving rise to the illusion of binaural beats. Cortical potentials recorded to binaural beats are distinct from onset responses. SIGNIFICANCE: Brain activity corresponding to an auditory illusion of low frequency beats can be recorded from the scalp.

N100 cortical potentials accompanying disrupted auditory nerve activity in auditory neuropathy (AN): effects of signal intensity and continuous noise.

OBJECTIVE: Auditory temporal processes in quiet are impaired in auditory neuropathy (AN) similar to normal hearing subjects tested in noise. N100 latencies were measured from AN subjects at several tone intensities in quiet and noise for comparison with a group of normal hearing individuals. METHODS: Subjects were tested with brief 100 ms tones (1.0 kHz, 100-40 dB SPL) in quiet and in continuous noise (90 dB SPL). N100 latency and amplitude were analyzed as a function of signal intensity and audibility. RESULTS: N100 latency in AN in quiet was delayed and amplitude was reduced compared to the normal group; the extent of latency delay was related to psychoacoustic measures of gap detection threshold and speech recognition scores, but not to audibility. Noise in normal hearing subjects was accompanied by N100 latency delays and amplitude reductions paralleling those found in AN tested in quiet. Additional N100 latency delays and amplitude reductions occurred in AN with noise. CONCLUSIONS: N100 latency to tones and performance on auditory temporal tasks were related in AN subjects. Noise masking in normal hearing subjects affected N100 latency to resemble AN in quiet. SIGNIFICANCE: N100 latency to tones may serve as an objective measure of the efficiency of auditory temporal processes.

Auditory-evoked potentials to frequency increase and decrease of high- and low-frequency tones.

OBJECTIVE: To define cortical brain responses to large and small frequency changes (increase and decrease) of high- and low-frequency tones. METHODS: Event-Related Potentials (ERPs) were recorded in response to a 10% or a 50% frequency increase from 250 or 4000 Hz tones that were approximately 3 s in duration and presented at 500-ms intervals. Frequency increase was followed after 1 s by a decrease back to base frequency. Frequency changes occurred at least 1 s before or after tone onset or offset, respectively. Subjects were not attending to the stimuli. Latency, amplitude and source current density estimates of ERPs were compared across frequency changes. RESULTS: All frequency changes evoked components P(50), N(100), and P(200). N(100) and P(200) had double peaks at bilateral and right temporal sites, respectively. These components were followed by a slow negativity (SN). The constituents of N(100) were predominantly localized to temporo-parietal auditory areas. The potentials and their intracranial distributions were affected by both base frequency (larger potentials to low frequency) and direction of change (larger potentials to increase than decrease), as well as by change magnitude (larger potentials to larger change). The differences between frequency increase and decrease depended on base frequency (smaller difference to high frequency) and were localized to frontal areas. CONCLUSIONS: Brain activity varies according to frequency change direction and magnitude as well as base frequency. SIGNIFICANCE: The effects of base frequency and direction of change may reflect brain networks involved in more complex processing such as speech that are differentially sensitive to frequency modulations of high (consonant discrimination) and low (vowels and prosody) frequencies.

Intensity changes in a continuous tone: auditory cortical potentials comparison with frequency changes.

OBJECTIVES: To examine auditory cortical potentials in normal-hearing subjects to intensity increments in a continuous pure tone at low, mid, and high frequency. METHODS: Electrical scalp potentials were recorded in response to randomly occurring 100 ms intensity increments of continuous 250, 1000, and 4000 Hz tones every 1.4 s. The magnitude of intensity change varied between 0, 2, 4, 6, and 8 dB above the 80 dB SPL continuous tone. RESULTS: Potentials included N100, P200, and a slow negative (SN) wave. N100 latencies were delayed whereas amplitudes were not affected for 250 Hz compared to 1000 and 4000 Hz. Functions relating the magnitude of the intensity change and N100 latency/amplitude did not differ in their slope among the three frequencies. No consistent relationship between intensity increment and SN was observed. Cortical dipole sources for N100 did not differ in location or orientation between the three frequencies. CONCLUSIONS: The relationship between intensity increments and N100 latency/amplitude did not differ between tonal frequencies. A cortical tonotopic arrangement was not observed for intensity increments. Our results are in contrast to prior studies of brain activities to brief frequency changes showing cortical tonotopic organization. SIGNIFICANCE: These results suggest that intensity and frequency discrimination employ distinct central processes.

Frequency changes in a continuous tone: auditory cortical potentials.

OBJECTIVE: We examined auditory cortical potentials in normal hearing subjects to spectral changes in continuous low and high frequency pure tones. METHODS: Cortical potentials were recorded to increments of frequency from continuous 250 or 4000Hz tones. The magnitude of change was random and varied from 0% to 50% above the base frequency. RESULTS: Potentials consisted of N100, P200 and a slow negative wave (SN). N100 amplitude, latency and dipole magnitude with frequency increments were significantly greater for low compared to high frequencies. Dipole amplitudes were greater in the right than left hemisphere for both base frequencies. The SN amplitude to frequency changes between 4% and 50% was not significantly related to the magnitude of spectral change. CONCLUSIONS: Modulation of N100 amplitude and latency elicited by spectral change is more pronounced with low compared to high frequencies. SIGNIFICANCE: These data provide electrophysiological evidence that central processing of spectral changes in the cortex differs for low and high frequencies. Some of these differences may be related to both temporal- and spectral-based coding at the auditory periphery. Central representation of frequency change may be related to the different temporal windows of integration across frequencies.

Human auditory steady-state responses.

Steady-state evoked potentials can be recorded from the human scalp in response to auditory stimuli presented at rates between 1 and 200 Hz or by periodic modulations of the amplitude and/or frequency of a continuous tone. Responses can be objectively detected using frequency-based analyses. In waking subjects, the responses are particularly prominent at rates near 40 Hz. Responses evoked by more rapidly presented stimuli are less affected by changes in arousal and can be evoked by multiple simultaneous stimuli without significant loss of amplitude. Response amplitude increases as the depth of modulation or the intensity increases. The phase delay of the response increases as the intensity or the carrier frequency decreases. Auditory steady-state responses are generated throughout the auditory nervous system, with cortical regions contributing more than brainstem generators to responses at lower modulation frequencies. These responses are useful for objectively evaluating auditory thresholds, assessing suprathreshold hearing, and monitoring the state of arousal during anesthesia.