Rebecca E. Millman

Second-order modulation detection thresholds for pure-tone and narrow-band noise carriers

Modulation perception has typically been characterized by measuring detection thresholds for sinusoidally amplitude-modulated (SAM) signals. This study uses multicomponent modulations. Second-order temporal modulation transfer functions (TMTFs) measure detection thresholds for a sinusoidal modulation of the modulation waveform of a SAM signal [Lorenzi et al., J. Acoust. Soc. Am. 110, 1030-2038 (2001)]. The SAM signal therefore acts as a carrier stimulus of frequency fm, and sinusoidal modulation of the SAM signals modulation depth (at rate fm) generates two additional components in the modulation spectrum at fm - fm and fm + fm. There is no spectral energy at the envelope beat frequency fm in the modulation spectrum of the physical stimulus. In the present study, second-order TMTFs were measured for three listeners when fm was 16, 64, and 256 Hz. The carrier was either a 5-kHz pure tone or a narrow-band noise with center frequency and bandwidth of 5 kHz and 2 Hz, respectively. The narrow-band noise carrier was used to prevent listeners from detecting spectral energy at the beat frequency fm in the internal stimulis modulation spectrum. The results show that, for the 5-kHz pure-tone carrier, second-order TMTFs are nearly low pass in shape; the overall sensitivity and cutoff frequency measured on these second-order TMTFs increase when fm increases from 16 to 256 Hz. For the 2-Hz-wide narrow-band noise carrier, second-order TMTFs are nearly flat in shape for fm = 16 and 64 Hz, and they show a high-pass segment for fm = 256 Hz. These results suggest that detection of spectral energy at the envelope beat frequency contributes in part to the detection of second-order modulation. This is consistent with the idea that nonlinear mechanisms in the auditory pathway produce an audible distortion component at the envelope beat frequency in the internal modulation spectrum of the sounds.

The role of phase-locking to the temporal envelope of speech in auditory perception and speech intelligibility

The temporal envelope of speech is important for speech intelligibility. Entrainment of cortical oscillations to the speech temporal envelope is a putative mechanism underlying speech intelligibility. Here we used magnetoencephalography (MEG) to test the hypothesis that phase-locking to the speech temporal envelope is enhanced for intelligible compared with unintelligible speech sentences. Perceptual “pop-out” was used to change the percept of physically identical tone-vocoded speech sentences from unintelligible to intelligible. The use of pop-out dissociates changes in phase-locking to the speech temporal envelope arising from acoustical differences between un/intelligible speech from changes in speech intelligibility itself. Novel and bespoke whole-head beamforming analyses, based on significant cross-correlation between the temporal envelopes of the speech stimuli and phase-locked neural activity, were used to localize neural sources that track the speech temporal envelope of both intelligible and unintelligible speech. Location-of-interest analyses were carried out in a priori defined locations to measure the representation of the speech temporal envelope for both un/intelligible speech in both the time domain (cross-correlation) and frequency domain (coherence). Whole-brain beamforming analyses identified neural sources phase-locked to the temporal envelopes of both unintelligible and intelligible speech sentences. Crucially there was no difference in phase-locking to the temporal envelope of speech in the pop-out condition in either the whole-brain or location-of-interest analyses, demonstrating that phase-locking to the speech temporal envelope is not enhanced by linguistic information.

Early Activity in Broca's Area During Reading Reflects Fast Access to Articulatory Codes From Print

Prior evidence for early activity in Brocas area during reading may reflect fast access to articulatory codes in left inferior frontal gyrus pars opercularis (LIFGpo). We put this hypothesis to test using a benchmark for articulatory involvement in reading known as the masked onset priming effect (MOPE). In masked onset priming, briefly presented pronounceable strings of letters that share an initial phoneme with subsequently presented target words (e.g., gilp-GAME) facilitate word naming responses compared with unrelated primes (dilp-GAME). Crucially, these priming effects only occur when the task requires articulation (naming), and not when it requires lexical decisions. A standard explanation of masked onset priming is that it reflects fast computation of articulatory output codes from letter representations. We therefore predicted 1) that activity in left IFG pars opercularis would be modulated by masked onset priming, 2) that priming-related modulation in LIFGpo would immediately follow activity in occipital cortex, and 3) that this modulation would be greater for naming than for lexical decision. These predictions were confirmed in a magnetoencephalography (MEG) priming study. MOPEs emerged in left IFG at ∼100 ms posttarget onset, and the priming effects were more sustained when the task involved articulation.

Magnified neural envelope coding predicts deficits in speech perception in noise

Verbal communication in noisy backgrounds is challenging. Understanding speech in background noise that fluctuates in intensity over time is particularly difficult for hearing-impaired listeners with a sensorineural hearing loss (SNHL). The reduction in fast-acting cochlear compression associated with SNHL exaggerates the perceived fluctuations in intensity in amplitude-modulated sounds. SNHL-induced changes in the coding of amplitude-modulated sounds may have a detrimental effect on the ability of SNHL listeners to understand speech in the presence of modulated background noise. To date, direct evidence for a link between magnified envelope coding and deficits in speech identification in modulated noise has been absent. Here, magnetoencephalography was used to quantify the effects of SNHL on phase locking to the temporal envelope of modulated noise (envelope coding) in human auditory cortex. Our results show that SNHL enhances the amplitude of envelope coding in posteromedial auditory cortex, whereas it enhances the fidelity of envelope coding in posteromedial and posterolateral auditory cortex. This dissociation was more evident in the right hemisphere, demonstrating functional lateralization in enhanced envelope coding in SNHL listeners. However, enhanced envelope coding was not perceptually beneficial. Our results also show that both hearing thresholds and, to a lesser extent, magnified cortical envelope coding in left posteromedial auditory cortex predict speech identification in modulated background noise. We propose a framework in which magnified envelope coding in posteromedial auditory cortex disrupts the segregation of speech from background noise, leading to deficits in speech perception in modulated background noise. SIGNIFICANCE STATEMENT People with hearing loss struggle to follow conversations in noisy environments. Background noise that fluctuates in intensity over time poses a particular challenge. Using magnetoencephalography, we demonstrate anatomically distinct cortical representations of modulated noise in normal-hearing and hearing-impaired listeners. This work provides the first link among hearing thresholds, the amplitude of cortical representations of modulated sounds, and the ability to understand speech in modulated background noise. In light of previous work, we propose that magnified cortical representations of modulated sounds disrupt the separation of speech from modulated background noise in auditory cortex.

Oscillatory Dynamics Supporting Semantic Cognition: MEG Evidence for the Contribution of the Anterior Temporal Lobe Hub and Modality-Specific Spokes

The “hub and spoke model” of semantic representation suggests that the multimodal features of objects are drawn together by an anterior temporal lobe (ATL) “hub”, while modality-specific “spokes” capture perceptual/action features. However, relatively little is known about how these components are recruited through time to support object identification. We used magnetoencephalography to measure neural oscillations within left ATL, lateral fusiform cortex (FC) and central sulcus (CS) during word-picture matching at different levels of specificity (employing superordinate vs. specific labels) for different categories (manmade vs. animal). This allowed us to determine (i) when each site was sensitive to semantic category and (ii) whether this was modulated by task demands. In ATL, there were two phases of response: from around 100 ms post-stimulus there were phasic bursts of low gamma activity resulting in reductions in oscillatory power, relative to a baseline period, that were modulated by both category and specificity; this was followed by more sustained power decreases across frequency bands from 250 ms onwards. In the spokes, initial power increases were not stronger for specific identification, while later power decreases were stronger for specific-level identification in FC for animals and in CS for manmade objects (from around 150 ms and 200 ms, respectively). These data are inconsistent with a temporal sequence in which early sensory-motor activity is followed by later retrieval in ATL. Instead, knowledge emerges from the rapid recruitment of both hub and spokes, with early specificity and category effects in the ATL hub. The balance between these components depends on semantic category and task, with visual cortex playing a greater role in the fine-grained identification of animals and motor cortex contributing to the identification of tools.

Causal cortical dynamics of a predictive enhancement of speech intelligibility

&NA; Speech perception may be underpinned by a hierarchical cortical system, which attempts to match “external” incoming sensory inputs with “internal” top‐down predictions. Prior knowledge modulates internal predictions of an upcoming stimulus and exerts its effects in temporal and inferior frontal cortex. Here, we used source‐space magnetoencephalography (MEG) to study the spatiotemporal dynamics underpinning the integration of prior knowledge in the speech processing network. Prior knowledge was manipulated to i) increase the perceived intelligibility of speech sentences, and ii) dissociate the perceptual effects of changes in speech intelligibility from acoustical differences in speech stimuli. Cortical entrainment to the speech temporal envelope, which accounts for neural activity specifically related to sensory information, was affected by prior knowledge: This effect emerged early (˜50 ms) in left inferior frontal gyrus (IFG) and then (˜100 ms) in Heschls gyrus (HG), and was sustained until latencies of ˜250 ms. Directed transfer function (DTF) measures were used for estimating direct Granger causal relations between locations of interest. In line with the cortical entrainment result, this analysis indicated that prior knowledge enhanced top‐down connections from left IFG to all the left temporal areas of interest – namely HG, superior temporal sulcus (STS), and middle temporal gyrus (MTG). In addition, intelligible speech increased top‐down information flow between left STS and left HG, and increased bottom‐up flow in higher‐order temporal cortex, specifically between STS and MTG. These results are compatible with theories that explain this mechanism as a result of both ascending and descending cortical interactions, such as predictive coding. Altogether, this study provides a detailed view of how, where and when prior knowledge influences continuous speech perception. HighlightsCortical entrainment to the speech envelope is modulated by prior knowledge.Prior knowledge enhances delta‐band entrainment.Enhanced envelope tracking in left IFG precedes the same effect in left HG.Intelligible speech modulates causal cortico‐cortical dynamics between temporal areas.

Effects of noise exposure on young adults with normal audiograms II: Behavioral measures

ABSTRACT An estimate of lifetime noise exposure was used as the primary predictor of performance on a range of behavioral tasks: frequency and intensity difference limens, amplitude modulation detection, interaural phase discrimination, the digit triplet speech test, the co‐ordinate response speech measure, an auditory localization task, a musical consonance task and a subjective report of hearing ability. One hundred and thirty‐eight participants (81 females) aged 18–36 years were tested, with a wide range of self‐reported noise exposure. All had normal pure‐tone audiograms up to 8 kHz. It was predicted that increased lifetime noise exposure, which we assume to be concordant with noise‐induced cochlear synaptopathy, would elevate behavioral thresholds, in particular for stimuli with high levels in a high spectral region. However, the results showed little effect of noise exposure on performance. There were a number of weak relations with noise exposure across the test battery, although many of these were in the opposite direction to the predictions, and none were statistically significant after correction for multiple comparisons. There were also no strong correlations between electrophysiological measures of synaptopathy published previously and the behavioral measures reported here. Consistent with our previous electrophysiological results, the present results provide no evidence that noise exposure is related to significant perceptual deficits in young listeners with normal audiometric hearing. It is possible that the effects of noise‐induced cochlear synaptopathy are only measurable in humans with extreme noise exposures, and that these effects always co‐occur with a loss of audiometric sensitivity. HIGHLIGHTSLarge study on the effects of lifetime noise exposure in normal‐hearing young adults.Performance on a range of behavioral tasks unrelated to noise exposure history.Effects of cochlear synaptopathy not evident in young audiometrically normal cohort.

Dynamic semantic cognition: Characterising coherent and controlled conceptual retrieval through time using magnetoencephalography and chronometric transcranial magnetic stimulation

Distinct neural processes are thought to support the retrieval of semantic information that is (i) coherent with strongly-encoded aspects of knowledge, and (ii) non-dominant yet relevant for the current task or context. While the brain regions that support readily coherent and more controlled patterns of semantic retrieval are relatively well-characterised, the temporal dynamics of these processes are not well-understood. This study used magnetoencephalography (MEG) and dual-pulse chronometric transcranial magnetic stimulation (cTMS) in two separate experiments to examine temporal dynamics during the retrieval of strong and weak associations. MEG results revealed a dissociation within left temporal cortex: anterior temporal lobe (ATL) showed greater oscillatory response for strong than weak associations, while posterior middle temporal gyrus (pMTG) showed the reverse pattern. Left inferior frontal gyrus (IFG), a site associated with semantic control and retrieval, showed both patterns at different time points. In the cTMS experiment, stimulation of ATL at ∼150 msec disrupted the efficient retrieval of strong associations, indicating a necessary role for ATL in coherent conceptual activations. Stimulation of pMTG at the onset of the second word disrupted the retrieval of weak associations, suggesting this site may maintain information about semantic context from the first word, allowing efficient engagement of semantic control. Together these studies provide converging evidence for a functional dissociation within the temporal lobe, across both tasks and time.

Task-based and resting-state fMRI reveal compensatory network changes following damage to left inferior frontal gyrus

Damage to left inferior prefrontal cortex in stroke aphasia is associated with semantic deficits reflecting poor control over conceptual retrieval, as opposed to loss of knowledge. However, little is known about how functional recruitment within the semantic network changes in patients with executive-semantic deficits. The current study acquired functional magnetic resonance imaging (fMRI) data from 14 patients with semantic aphasia, who had difficulty with flexible semantic retrieval following left prefrontal damage, and 16 healthy age-matched controls, allowing us to examine activation and connectivity in the semantic network. We examined neural activity while participants listened to spoken sentences that varied in their levels of lexical ambiguity and during rest. We found group differences in two regions thought to be good candidates for functional compensation: ventral anterior temporal lobe (vATL), which is strongly implicated in comprehension, and posterior middle temporal gyrus (pMTG), which is hypothesized to work together with left inferior prefrontal cortex to support controlled aspects of semantic retrieval. The patients recruited both of these sites more than controls in response to meaningful sentences. Subsequent analysis identified that, in control participants, the recruitment of pMTG to ambiguous sentences was inversely related to functional coupling between pMTG and anterior superior temporal gyrus (aSTG) at rest, while the patients showed the opposite pattern. Moreover, stronger connectivity between pMTG and aSTG in patients was associated with better performance on a test of verbal semantic association, suggesting that this temporal lobe connection supports comprehension in the face of damage to left inferior prefrontal cortex. These results characterize network changes in patients with executive-semantic deficits and converge with studies of healthy participants in providing evidence for a distributed system underpinning semantic control that includes pMTG in addition to left inferior prefrontal cortex.

Supra-threshold auditory brainstem response amplitudes in humans: Test-retest reliability, electrode montage and noise exposure

&NA; The auditory brainstem response (ABR) is a sub‐cortical evoked potential in which a series of well‐defined waves occur in the first 10 ms after the onset of an auditory stimulus. Wave V of the ABR, particularly wave V latency, has been shown to be remarkably stable over time in individual listeners. However, little attention has been paid to the reliability of wave I, which reflects auditory nerve activity. This ABR component has attracted interest recently, as wave I amplitude has been identified as a possible non‐invasive measure of noise‐induced cochlear synaptopathy. The current study aimed to determine whether ABR wave I amplitude has sufficient test‐retest reliability to detect impaired auditory nerve function in an otherwise normal‐hearing listener. Thirty normal‐hearing females were tested, divided equally into low‐ and high‐noise exposure groups. The stimulus was an 80 dB nHL click. ABR recordings were made from the ipsilateral mastoid and from the ear canal (using a tiptrode). Although there was some variability between listeners, wave I amplitude had high test‐retest reliability, with an intraclass correlation coefficient (ICC) comparable to that for wave V amplitude. There were slight gains in reliability for wave I amplitude when recording from the ear canal (ICC of 0.88) compared to the mastoid (ICC of 0.85). The summating potential (SP) and ratio of SP to wave I were also quantified and found to be much less reliable than measures of wave I and V amplitude. Finally, we found no significant differences in the amplitude of any wave components between low‐ and high‐noise exposure groups. We conclude that, if the other sources of between‐subject variability can be controlled, wave I amplitude is sufficiently reliable to accurately characterize individual differences in auditory nerve function. HighlightsABR wave I and V amplitudes have excellent test‐retest reliability in humans.SP amplitude and SP/AP ratio have poor test‐retest reliability.Canal tiptrodes result in only slightly increased reliability re. mastoid electrodes.No significant differences in amplitudes between low‐ and high‐noise exposed females.