Amy R. Horwitz

A potential role for thalamocingulate circuitry in human maternal behavior

BACKGROUND Little is known about the regional brain basis of human maternal behavior. To understand this better, we have been examining brain activity in mothers listening to infant cries. METHODS We measured functional Magnetic Resonance Imaging brain activity in healthy, breastfeeding first-time mothers with young infants while they listened to infant cries, white noise control sounds, and a rest condition. Based on the thalamocingulate theory of maternal behavior and pilot work, we hypothesized that the cingulate, medial thalamus, medial prefrontal cortex, and right orbitofrontal cortex would display more activity with infant cries than with white noise (comparison 1) and would uniquely activate with the cries, meaning that these regions would display activity with cry minus rest but not with white noise minus rest (comparison 2). RESULTS In hypothesized regions, the group displayed more activity in the medial thalamus, medial prefrontal and right orbitofrontal cortices with both comparisons. The anterior and posterior cingulate cortex displayed more activity only with comparison 1. In non-hypothesized brain regions, several other structures thought important in rodent maternal behavior displayed activity with both comparisons including the midbrain, hypothalamus, dorsal and ventral striatum, and vicinity of the lateral septal region. CONCLUSIONS Our results partially support our hypotheses and are generally consistent with neuroanatomical studies of rodent maternal behavior.

At the heart of the ventral attention system: The right anterior insula

The anterior insula has been hypothesized to provide a link between attention‐related problem solving and salience systems during the coordination and evaluation of task performance. Here, we test the hypothesis that the anterior insula/medial frontal operculum (aI/fO) provides linkage across systems supporting task demands and attention systems by examining the patterns of functional connectivity during word recognition and spatial attention functional imaging tasks. A shared set of frontal regions (right aI/fO, right dorsolateral prefrontal cortex, bilateral anterior cingulate) were engaged, regardless of perceptual domain (auditory or visual) or mode of response (word production or button press). We present novel evidence that: (1) the right aI/fO is functionally connected with other frontal regions implicated in executive function and not just brain regions responsive to stimulus salience; and (2) that the aI/fO, but not the ACC, exhibits significantly correlated activity with other brain regions specifically engaged by tasks with varying perceptual and behavioral demands. These results support the hypothesis that the right aI/fO aids in the coordination and evaluation of task performance across behavioral tasks with varying perceptual and response demands. Hum Brain Mapp 2009.

Recovery from prior stimulation: masking of speech by interrupted noise for younger and older adults with normal hearing.

In a previous study [Dubno et al, J. Acoust. Soc. Am. 111, 2897-2907 (2002)], older subjects benefitted less than younger subjects from momentary improvements in signal-to-noise ratio when listening to speech in interrupted maskers. It has been hypothesized that the benefit derived from interrupted maskers may be related to recovery from forward masking, i.e., the recovery of a response to a suprathreshold signal from prior stimulation by a masker. The effect of interrupted maskers on speech recognition may be well suited to test hypotheses regarding recovery from prior stimulation, given that both involve the perception of signals following a masker. Here, younger and older adults with normal but not identical audiograms listened to nonsense syllables at moderate and high levels in a speech-shaped noise that was modulated by a 2-, 10-, 25-, or 50-Hz square wave. An additional low-level noise was always present that was shaped to produce equivalent masked thresholds for all subjects. To assess recovery from forward masking, forward-masked thresholds were measured at 0.5 and 4.0 kHz as a function of the delay between the speech-shaped masker and the signal. Speech recognition in interrupted noise was poorer for older than younger subjects. Small but consistent age-related differences were observed in the decrease in score with interrupted noise relative to the score without interrupted noise. Forward-masked thresholds of older subjects were higher than those of younger subjects, but there were no age-related differences in the amount of forward masking or in simultaneous masking. Negative correlations were observed between speech-recognition scores in interrupted noise and forward-masked thresholds. That is, the benefit derived from momentary improvements in speech audibility in an interrupted noise decreased as forward-masked thresholds increased. Stronger correlations with forward masking were observed for the higher frequency signal, for higher noise interruption rates, and when the signal-to-noise ratio was poor. Comparisons of speech-recognition scores at moderate and high levels for younger and older subjects were not consistent with the hypothesis of an age-related difference in the contribution of low-spontaneous-rate fibers to speech recognition in interrupted noise.

Use of context by young and aged adults with normal hearing

Word recognition in sentences with and without context was measured in young and aged subjects with normal but not identical audiograms. Benefit derived from context by older adults has been obscured, in part, by the confounding effect of even mildly elevated thresholds, especially as listening conditions vary in difficulty. This problem was addressed here by precisely controlling signal-to-noise ratio across conditions and by accounting for individual differences in signal-to-noise ratio. Pure-tone thresholds and word recognition were measured in quiet and threshold-shaped maskers that shifted quiet thresholds by 20 and 40 dB. Word recognition was measured at several speech levels in each condition. Threshold was defined as the speech level (or signal-to-noise ratio) corresponding to the 50 rau point on the psychometric function. As expected, thresholds and slopes of psychometric functions were different for sentences with context compared to those for sentences without context. These differences were equivalent for young and aged subjects. Individual differences in word recognition among all subjects, young and aged, were accounted for by individual differences in signal-to-noise ratio. With signal-to-noise ratio held constant, word recognition for all subjects remained constant or decreased only slightly as speech and noise levels increased. These results suggest that, given equivalent speech audibility, older and younger listeners derive equivalent benefit from context.

The Time Course of Hearing Aid Benefit

Objective: The present study sought to determine whether benefit derived from hearing aid use increases with hearing aid experience. This question is of considerable interest to both researchers and clinicians. Several previous investigations recently published have obtained mixed results. Design: Control conditions were provided to address the potential influences of practice effects and changes in preferred hearing aid gain. The experimental group consisted of 13 hearing‐impaired listeners about to be fit with their first hearing aid. The control group consisted of 13 hearing aid users with at least 1 yrs experience with their hearing aids. The control group permitted examination of practice effects that may have confounded previous results showing increases in benefit with experience. Hearing aid benefit was defined as aided speech recognition ability minus unaided speech recognition ability and was assessed repeatedly over 18 wk. Two measures of hearing aid benefit were employed: an objective syllable recognition task and a subjective questionnaire. For the objective measure, hearing aid benefit was assessed for the condition of fixed hearing aid gain and also for the condition of subject‐adjusted hearing aid gain to examine effects of changes in audibility that may have influenced benefit and confounded previous results. Results: The objective measure of group mean hearing aid benefit increased significantly over time for both gain conditions for the new hearing aid users, but did not increase for the long‐standing control group. Subjective benefit increased over time, but without statistical significance for the new hearing aid users, and was essentially unchanged for the long‐standing control group. Conclusions: Results suggest that the observed improvements in speech recognition are not due to increases in audibility nor to simple practice effects. The overall improvements in benefit over time were of statistical significance and also practical importance for studies of group differences. However, the improvements are too small to be observed consistently for individual hearing aid users.

Longitudinal changes in speech recognition in older persons

Recognition of isolated monosyllabic words in quiet and recognition of key words in low- and high-context sentences in babble were measured in a large sample of older persons enrolled in a longitudinal study of age-related hearing loss. Repeated measures were obtained yearly or every 2 to 3 years. To control for concurrent changes in pure-tone thresholds and speech levels, speech-recognition scores were adjusted using an importance-weighted speech-audibility metric (AI). Linear-regression slope estimated the rate of change in adjusted speech-recognition scores. Recognition of words in quiet declined significantly faster with age than predicted by declines in speech audibility. As subjects aged, observed scores deviated increasingly from AI-predicted scores, but this effect did not accelerate with age. Rate of decline in word recognition was significantly faster for females than males and for females with high serum progesterone levels, whereas noise history had no effect. Rate of decline did not accelerate with age but increased with degree of hearing loss, suggesting that with more severe injury to the auditory system, impairments to auditory function other than reduced audibility resulted in faster declines in word recognition as subjects aged. Recognition of key words in low- and high-context sentences in babble did not decline significantly with age.

Level-dependent changes in detection of temporal gaps in noise markers by adults with normal and impaired hearing

Compression in the basilar-membrane input-output response flattens the temporal envelope of a fluctuating signal when more gain is applied to lower level than higher level temporal components. As a result, level-dependent changes in gap detection for signals with different depths of envelope fluctuation and for subjects with normal and impaired hearing may reveal effects of compression. To test these assumptions, gap detection with and without a broadband noise was measured with 1, 000-Hz-wide (flatter) and 50-Hz-wide (fluctuating) noise markers as a function of marker level. As marker level increased, background level also increased, maintaining a fixed acoustic signal-to-noise ratio (SNR) to minimize sensation-level effects on gap detection. Significant level-dependent changes in gap detection were observed, consistent with effects of cochlear compression. For the flatter marker, gap detection that declines with increases in level up to mid levels and improves with further increases in level may be explained by an effective flattening of the temporal envelope at mid levels, where compression effects are expected to be strongest. A flatter effective temporal envelope corresponds to a reduced effective SNR. The effects of a reduction in compression (resulting in larger effective SNRs) may contribute to better-than-normal gap detection observed for some hearing-impaired listeners.

Estimates of basilar-membrane nonlinearity effects on masking of tones and speech.

Objective: The aim of this experiment was to assess the contribution of cochlear nonlinearities to speech recognition in noise for individuals with normal hearing and a range of quiet thresholds. For signals close to the characteristic frequency (CF) of a place on the basilar membrane, the normal growth of response of the basilar membrane is linear at lower signal levels and compressed at medium to higher signal levels. In contrast, at moderate to high CFs, the basilar membrane responds more linearly to stimuli at frequencies well below the CF regardless of input level. Thus, for moderate-level speech and a lower frequency masker, the response to the masker grows linearly whereas the response to the speech is compressed, which may result in changes in the effectiveness of the masker on speech recognition with increases in masker level. To test this hypothesis, observed speech-recognition scores were compared with scores predicted using an audibility-based model, which did not include nonlinear effects that may influence masker effectiveness. Design: Growth of simultaneous masking was measured for moderate-level bandpass-filtered nonsense syllables and for 350-msec pure tones at frequencies within the speech passband. Masker frequencies were within (on-frequency) or below (off-frequency) the speech passband. Estimates of basilar-membrane nonlinearities were derived from growth-of-masking functions for 10-msec, 2.0- and 4.0-kHz tones in narrowband, off-frequency maskers presented simultaneously. Subjects were 26 adults with normal hearing with approximately a 20-dB range of average quiet thresholds. Results: Breakpoints (i.e., the levels corresponding to the transitions from linear to nonlinear responses) were strongly associated with quiet thresholds but slopes measured above the breakpoints were independent of quiet thresholds. Individual differences were substantially larger for off-frequency masking of pure tones and speech than for on-frequency masking of pure tones and speech. Using an audibility-based predictive model, the change in speech audibility resulting from the compressed response to speech with increasing off-frequency masker level (and the resulting decline in scores) was well predicted from nonlinear growth of masking for pure tones measured in the same off-frequency masker. However, absolute speech-recognition predictions were generally inaccurate and were a function of how well pure-tone signal levels at masked threshold estimated masker effectiveness for speech. That is, subjects with lower off-frequency masked thresholds had less accurate predictions of speech recognition in off-frequency maskers. Conclusions: Large individual differences in off-frequency masking of pure tones and speech are consistent with the assumption that small changes in the shape of the basilar-membrane input-output function result in large changes in the amount of off-frequency masking but small (if any) changes in on-frequency masking where the signal and masker are subject to a similar compression. Growth of off-frequency masking of pure tones and speech were correlated with each other, consistent with the underlying basilar-membrane response, and consistent with changes in breakpoints for subjects with normal hearing and a range of quiet thresholds. These results provide support for a role of nonlinear effects in the understanding of speech in noise.

Individual Differences in Behavioral Estimates of Cochlear Nonlinearities

Psychophysical methods provide a mechanism to infer the characteristics of basilar membrane responses in humans that cannot be directly measured. Because these behavioral measures are indirect, the interpretation of results depends on several underlying assumptions. Ongoing uncertainty about the suitability of these assumptions and the most appropriate measurement and compression estimation procedures, and unanswered questions regarding the effects of cochlear hearing loss and age on basilar membrane nonlinearities, motivated this experiment. Here, estimates of cochlear nonlinearities using temporal masking curves (TMCs) were obtained in a large sample of adults of various ages whose hearing ranged from normal to moderate cochlear hearing loss (Experiment 1). A wide range of compression slopes was observed, even for subjects with similar ages and thresholds, which warranted further investigation (Experiment 2). Potential sources of variance contributing to these individual differences were explored, including procedural-related factors (test–retest reliability, suitability of the linear-reference TMC, probe sensation levels, and parameters of TMC fitting algorithms) and subject-related factors (age and age-related changes in temporal processing, strength of cochlear nonlinearities estimated with distortion-product otoacoustic emissions, estimates of changes in cochlear function from damage to outer hair cells versus inner hair cells). Subject age did not contribute significantly to TMC or compression slopes, and TMC slopes did not vary significantly with threshold. Test–retest reliability of TMCs suggested that TMC masker levels and the general shapes of TMCs did not change in a systematic way when re-measured many weeks later. Although the strength of compression decreased slightly with increasing hearing loss, the magnitude of individual differences in compression estimates makes it difficult to determine the effects of hearing loss and cochlear damage on basilar membrane nonlinearities in humans.

Individual and level-dependent differences in masking for adults with normal and impaired hearing

Simultaneous, on-frequency masking is commonly assumed to be linear with increasing noise intensity. However, some evidence suggests that, expressed in terms of signal-to-noise ratio changes with background level changes, masking slopes can vary from 0 dB/dB. These results and evidence from a large sample of subjects with normal and impaired hearing demonstrate level-dependent changes in masking, large individual differences in masking among subjects with similar thresholds in quiet, and significant correlations of masking slope with other estimates of auditory function measured in the same backgrounds.