Modulation Rate Discrimination Research Articles

Population coding of time-varying sounds in the nonlemniscal inferior colliculus.

The inferior colliculus (IC) of the midbrain is important for complex sound processing, such as discriminating conspecific vocalizations and human speech. The IC's nonlemniscal, dorsal "shell" region is likely important for this process, as neurons in these layers project to higher-order thalamic nuclei that subsequently funnel acoustic signals to the amygdala and nonprimary auditory cortices, forebrain circuits important for vocalization coding in a variety of mammals, including humans. However, the extent to which shell IC neurons transmit acoustic features necessary to discern vocalizations is less clear, owing to the technical difficulty of recording from neurons in the IC's superficial layers via traditional approaches. Here, we use two-photon Ca2+ imaging in mice of either sex to test how shell IC neuron populations encode the rate and depth of amplitude modulation, important sound cues for speech perception. Most shell IC neurons were broadly tuned, with a low neurometric discrimination of amplitude modulation rate; only a subset was highly selective to specific modulation rates. Nevertheless, neural network classifier trained on fluorescence data from shell IC neuron populations accurately classified amplitude modulation rate, and decoding accuracy was only marginally reduced when highly tuned neurons were omitted from training data. Rather, classifier accuracy increased monotonically with the modulation depth of the training data, such that classifiers trained on full-depth modulated sounds had median decoding errors of ∼0.2 octaves. Thus, shell IC neurons may transmit time-varying signals via a population code, with perhaps limited reliance on the discriminative capacity of any individual neuron.NEW & NOTEWORTHY The IC's shell layers originate a "nonlemniscal" pathway important for perceiving vocalization sounds. However, prior studies suggest that individual shell IC neurons are broadly tuned and have high response thresholds, implying a limited reliability of efferent signals. Using Ca2+ imaging, we show that amplitude modulation is accurately represented in the population activity of shell IC neurons. Thus, downstream targets can read out sounds' temporal envelopes from distributed rate codes transmitted by populations of broadly tuned neurons.

Auditory cortex plasticity supports social learning

The acquisition of new auditory skills can be facilitated by experiencing a conspecific performing a well-defined behavior (i.e., social learning). Although the neural bases for auditory social learning remain uncertain, one plausible hypothesis is that social experience induces long-term changes to auditory cortex, thereby facilitating the subsequent acquisition of an auditory skill. To explore this idea, we developed a social learning paradigm in which naïve observer gerbils are exposed to a demonstrator that performs an amplitude modulation (AM) rate discrimination task on the other side of an opaque divider. Thus, observers have no access to visual cues. When exposed to a demonstrator for five successive days, observers subsequently acquire the AM task more rapidly than controls. Two experiments suggest that auditory cortex is necessary and sufficient for social learning. First, inactivating the observer’s auditory cortex during each social exposure session significantly delayed task acquisition. Second, recordings from the observer’s auditory cortex revealed that individual neurons displayed a significant improvement in AM stimulus discrimination across the five exposure sessions, and the magnitude of neural improvement correlated with an animal’s subsequent rate of task acquisition. Together, these findings suggest that auditory cortex plasticity plays a pivotal role in social learning.

Relationships Between the Auditory Nerve Sensitivity to Amplitude Modulation, Perceptual Amplitude Modulation Rate Discrimination Sensitivity, and Speech Perception Performance in Postlingually Deafened Adult Cochlear Implant Users.

This study assessed the relationships between the salience of amplitude modulation (AM) cues encoded at the auditory nerve (AN), perceptual sensitivity to changes in AM rate (i.e., AM rate discrimination threshold, AMRDT), and speech perception scores in postlingually deafened adult cochlear implant (CI) users. Study participants were 18 postlingually deafened adults with Cochlear Nucleus devices, including five bilaterally implanted patients. For each of 23 implanted ears, neural encoding of AM cues at 20 Hz at the AN was evaluated at seven electrode locations across the electrode array using electrophysiological measures of the electrically evoked compound action potential (eCAP). The salience of AM neural encoding was quantified by the Modulated Response Amplitude Ratio (MRAR). Psychophysical measures of AMRDT for 20 Hz modulation were evaluated in 16 ears using a three-alternative, forced-choice procedure, targeting 79.4% correct on the psychometric function. AMRDT was measured at up to five electrode locations for each test ear, including the electrode pair that showed the largest difference in the MRAR. Consonant-Nucleus-Consonant (CNC) word scores presented in quiet and in speech-shaped noise at a signal to noise ratio (SNR) of +10 dB were measured in all 23 implanted ears. Simulation tests were used to assess the variations in correlation results when using the MRAR and AMRDT measured at only one electrode location in each participant to correlate with CNC word scores. Linear Mixed Models (LMMs) were used to evaluate the relationship between MRARs/AMRDTs measured at individual electrode locations and CNC word scores. Spearman Rank correlation tests were used to evaluate the strength of association between CNC word scores measured in quiet and in noise with (1) the variances in MRARs and AMRDTs, and (2) the averaged MRAR or AMRDT across multiple electrodes tested for each participant. There was no association between the MRAR and AMRDT. Using the MRAR and AMRDT measured at only one, randomly selected electrode location to assess their associations with CNC word scores could lead to opposite conclusions. Both the results of LMMs and Spearman Rank correlation tests showed that CNC word scores measured in quiet or at 10 dB SNR were not significantly correlated with the MRAR or AMRDT. In addition, the results of Spearman Rank correlation tests showed that the variances in MRARs and AMRDTs were not significantly correlated with CNC word scores measured in quiet or in noise. The difference in AN sensitivity to AM cues is not the primary factor accounting for the variation in AMRDTs measured at different stimulation sites within individual CI users. The AN sensitivity to AM per se may not be a crucial factor for CNC word perception in quiet or at 10 dB SNR in postlingually deafened adult CI users. Using electrophysiological or psychophysical results measured at only one electrode location to correlate with speech perception scores in CI users can lead to inaccurate, if not wrong, conclusions.

Memory Specific to Temporal Features of Sound Is Formed by Cue-Selective Enhancements in Temporal Coding Enabled by Inhibition of an Epigenetic Regulator

Recent investigation of memory-related functions in the auditory system have capitalized on the use of memory-modulating molecules to probe the relationship between memory and substrates of memory in auditory system coding. For example, epigenetic mechanisms, which regulate gene expression necessary for memory consolidation, are powerful modulators of learning-induced neuroplasticity and long-term memory (LTM) formation. Inhibition of the epigenetic regulator histone deacetylase 3 (HDAC3) promotes LTM, which is highly specific for spectral features of sound. The present work demonstrates for the first time that HDAC3 inhibition also enables memory for temporal features of sound. Adult male rats trained in an amplitude modulation (AM) rate discrimination task and treated with a selective inhibitor of HDAC3 formed memory that was highly specific to the AM rate paired with reward. Sound-specific memory revealed behaviorally was associated with a signal-specific enhancement in temporal coding in the auditory system; stronger phase locking that was specific to the rewarded AM rate was revealed in both the surface-recorded frequency following response and auditory cortical multiunit activity in rats treated with the HDAC3 inhibitor. Furthermore, HDAC3 inhibition increased trial-to-trial cortical response consistency (relative to naive and trained vehicle-treated rats), which generalized across different AM rates. Stronger signal-specific phase locking correlated with individual behavioral differences in memory specificity for the AM signal. These findings support that epigenetic mechanisms regulate activity-dependent processes that enhance discriminability of sensory cues encoded into LTM in both spectral and temporal domains, which may be important for remembering spectrotemporal features of sounds, for example, as in human voices and speech.SIGNIFICANCE STATEMENTEpigenetic mechanisms have recently been implicated in memory and information processing. Here, we use a pharmacological inhibitor of HDAC3 in a sensory model of learning to reveal the ability of HDAC3 to enable precise memory for amplitude-modulated sound cues. In so doing, we uncover neural substrates for memory's specificity for temporal sound cues. Memory specificity was supported by auditory cortical changes in temporal coding, including greater response consistency and stronger phase locking. HDAC3 appears to regulate effects across domains that determine specific cue saliency for behavior. Thus, epigenetic players may gate how sensory information is stored in long-term memory and can be leveraged to reveal the neural substrates of sensory details stored in memory.

Easy-to-hard effects in perceptual learning depend upon the degree to which initial trials are "easy".

Starting perceptual training at easy levels before progressing to difficult levels generally produces better learning outcomes than constantly difficult training does. However, little is known about how "easy" these initial levels should be in order to yield easy-to-hard effects. We compared five levels of initial training block difficulty varying from very easy to hard in two auditory-discrimination learning tasks-a frequency modulation rate discrimination (Experiment 1) and a frequency range discrimination (Experiment 2). The degree of difficulty was based on individualized pretraining ~71% correct discrimination thresholds. Both experiments revealed a sweet spot for easy-to-hard effects. Conditions where initial blocks were either too easy or too difficult produced less benefit than did blocks of intermediate difficulty. Results challenge assumptions that sequencing effects in learning are related to attentional spotlighting of task-relevant dimensions. Rather, they support incremental learning models that account for easy-to-hard effects. Further, the results have implications for how perceptual training regimens should be designed to maximize the benefits of rehabilitative perceptual training.

Temporal Sensitivity Measured Shortly After Cochlear Implantation Predicts 6-Month Speech Recognition Outcome.

Psychoacoustic tests assessed shortly after cochlear implantation are useful predictors of the rehabilitative speech outcome. While largely independent, both spectral and temporal resolution tests are important to provide an accurate prediction of speech recognition. However, rapid tests of temporal sensitivity are currently lacking. Here, we propose a simple amplitude modulation rate discrimination (AMRD) paradigm that is validated by predicting future speech recognition in adult cochlear implant (CI) patients. In 34 newly implanted patients, we used an adaptive AMRD paradigm, where broadband noise was modulated at the speech-relevant rate of ~4 Hz. In a longitudinal study, speech recognition in quiet was assessed using the closed-set Freiburger number test shortly after cochlear implantation (t0) as well as the open-set Freiburger monosyllabic word test 6 months later (t6). Both AMRD thresholds at t0 (r = -0.51) and speech recognition scores at t0 (r = 0.56) predicted speech recognition scores at t6. However, AMRD and speech recognition at t0 were uncorrelated, suggesting that those measures capture partially distinct perceptual abilities. A multiple regression model predicting 6-month speech recognition outcome with deafness duration and speech recognition at t0 improved from adjusted R = 0.30 to adjusted R = 0.44 when AMRD threshold was added as a predictor. These findings identify AMRD thresholds as a reliable, nonredundant predictor above and beyond established speech tests for CI outcome. This AMRD test could potentially be developed into a rapid clinical temporal-resolution test to be integrated into the postoperative test battery to improve the reliability of speech outcome prognosis.

Developmental hearing loss impedes auditory task learning and performance in gerbils

The consequences of developmental hearing loss have been reported to include both sensory and cognitive deficits. To investigate these issues in a non-human model, auditory learning and asymptotic psychometric performance were compared between normal hearing (NH) adult gerbils and those reared with conductive hearing loss (CHL). At postnatal day 10, before ear canal opening, gerbil pups underwent bilateral malleus removal to induce a permanent CHL. Both CHL and control animals were trained to approach a water spout upon presentation of a target (Go stimuli), and withhold for foils (Nogo stimuli). To assess the rate of task acquisition and asymptotic performance, animals were tested on an amplitude modulation (AM) rate discrimination task. Behavioral performance was calculated using a signal detection theory framework. Animals reared with developmental CHL displayed a slower rate of task acquisition for AM discrimination task. Slower acquisition was explained by an impaired ability to generalize to newly introduced stimuli, as compared to controls. Measurement of discrimination thresholds across consecutive testing blocks revealed that CHL animals required a greater number of testing sessions to reach asymptotic threshold values, as compared to controls. However, with sufficient training, CHL animals approached control performance. These results indicate that a sensory impediment can delay auditory learning, and increase the risk of poor performance on a temporal task.

Effects of elevated amplitude modulation discrimination threshold on simultaneous amplitude modulation rate discrimination

Cochlear-implant (CI) users struggle to understand speech in difficult listening environments partly because they have limited access to auditory cues that allow for perceptual segregation between target and competing sounds. Because CI speech coding strategies optimize the encoding of envelope cues to promote better speech intelligibility, envelope may be an important cue for source segregation in CI users. Recent work has suggested that access to amplitude modulation (AM) depth and rate varies according to location of electrodes in the cochlea, and that turning off electrodes that yield poorer AM sensitivity may improve speech reception. The purpose of this study was to investigate the role of AM sensitivity in the ability of listeners to use AM rate information in multiple electrodes. It was hypothesized that electrodes with higher AM thresholds, which were simulated in normal-hearing listeners, would limit listeners’ ability to segregate AM rates. Subjects discriminated between pairs of stimuli that varied in AM rate relative to a reference rate. Stimulus pairing was within- or across-ears. Results suggest that AM insensitive electrodes impair CI users’ ability to discriminate AM rates, which may limit source segregation using envelope cues in complex listening environments. [This work was supported by NIH-NIDCD R01 DC003083 (Litovsky).]

Deficits in the pitch sensitivity of cochlear-implanted children speaking English or Mandarin.

Sensitivity to complex pitch is notoriously poor in adults with cochlear implants (CIs), but it is unclear whether this is true for children with CIs. Many are implanted today at a very young age, and factors related to brain plasticity (age at implantation, duration of CI experience, and speaking a tonal language) might have strong influences on pitch sensitivity. School-aged children participated, speaking English or Mandarin, having normal hearing (NH) or wearing a CI, using their clinically assigned settings with envelope-based coding strategies. Percent correct was measured in three-interval three-alternative forced choice tasks, for the discrimination of fundamental frequency (F0) of broadband harmonic complexes, and for the discrimination of sinusoidal amplitude modulation rate (AMR) of broadband noise, with reference frequencies at 100 and 200 Hz to focus on voice pitch processing. Data were fitted using a maximum-likelihood technique. CI children displayed higher thresholds and shallower slopes than NH children in F0 discrimination, regardless of linguistic background. Thresholds and slopes were more similar between NH and CI children in AMR discrimination. Once the effect of chronological age was extracted from the variance, the aforementioned factors related to brain plasticity did not contribute significantly to the CI children's sensitivity to pitch. Unless different strategies attempt to encode fine structure information, potential benefits of plasticity may be missed.

The Brain Dynamics of Rapid Perceptual Adaptation to Adverse Listening Conditions

Listeners show a remarkable ability to quickly adjust to degraded speech input. Here, we aimed to identify the neural mechanisms of such short-term perceptual adaptation. In a sparse-sampling, cardiac-gated functional magnetic resonance imaging (fMRI) acquisition, human listeners heard and repeated back 4-band-vocoded sentences (in which the temporal envelope of the acoustic signal is preserved, while spectral information is highly degraded). Clear-speech trials were included as baseline. An additional fMRI experiment on amplitude modulation rate discrimination quantified the convergence of neural mechanisms that subserve coping with challenging listening conditions for speech and non-speech. First, the degraded speech task revealed an "executive" network (comprising the anterior insula and anterior cingulate cortex), parts of which were also activated in the non-speech discrimination task. Second, trial-by-trial fluctuations in successful comprehension of degraded speech drove hemodynamic signal change in classic "language" areas (bilateral temporal cortices). Third, as listeners perceptually adapted to degraded speech, downregulation in a cortico-striato-thalamo-cortical circuit was observable. The present data highlight differential upregulation and downregulation in auditory-language and executive networks, respectively, with important subcortical contributions when successfully adapting to a challenging listening situation.

Navigating the Auditory Scene: An Expert Role for the Hippocampus

Over a typical career piano tuners spend tens of thousands of hours exploring a specialized acoustic environment. Tuning requires accurate perception and adjustment of beats in two-note chords that serve as a navigational device to move between points in previously learned acoustic scenes. It is a two-stage process that depends on the following: first, selective listening to beats within frequency windows, and, second, the subsequent use of those beats to navigate through a complex soundscape. The neuroanatomical substrates underlying brain specialization for such fundamental organization of sound scenes are unknown. Here, we demonstrate that professional piano tuners are significantly better than controls matched for age and musical ability on a psychophysical task simulating active listening to beats within frequency windows that is based on amplitude modulation rate discrimination. Tuners show a categorical increase in gray matter volume in the right frontal operculum and right superior temporal lobe. Tuners also show a striking enhancement of gray matter volume in the anterior hippocampus, parahippocampal gyrus, and superior temporal gyrus, and an increase in white matter volume in the posterior hippocampus as a function of years of tuning experience. The relationship with gray matter volume is sensitive to years of tuning experience and starting age but not actual age or level of musicality. Our findings support a role for a core set of regions in the hippocampus and superior temporal cortex in skilled exploration of complex sound scenes in which precise sound "templates" are encoded and consolidated into memory over time in an experience-dependent manner.

Auditory skills and brain morphology predict individual differences in adaptation to degraded speech

Noise-vocoded speech is a spectrally highly degraded signal, but it preserves the temporal envelope of speech. Listeners vary considerably in their ability to adapt to this degraded speech signal. Here, we hypothesised that individual differences in adaptation to vocoded speech should be predictable by non-speech auditory, cognitive, and neuroanatomical factors. We tested 18 normal-hearing participants in a short-term vocoded speech-learning paradigm (listening to 100 4-band-vocoded sentences). Non-speech auditory skills were assessed using amplitude modulation (AM) rate discrimination, where modulation rates were centred on the speech-relevant rate of 4Hz. Working memory capacities were evaluated (digit span and nonword repetition), and structural MRI scans were examined for anatomical predictors of vocoded speech learning using voxel-based morphometry. Listeners who learned faster to understand degraded speech also showed smaller thresholds in the AM discrimination task. This ability to adjust to degraded speech is furthermore reflected anatomically in increased grey matter volume in an area of the left thalamus (pulvinar) that is strongly connected to the auditory and prefrontal cortices. Thus, individual non-speech auditory skills and left thalamus grey matter volume can predict how quickly a listener adapts to degraded speech.

Sensitivity of school-aged children to pitch-related cues

Two experiments investigated the ability of 17 school-aged children to process purely temporal and spectro-temporal cues that signal changes in pitch. Percentage correct was measured for the discrimination of sinusoidal amplitude modulation rate (AMR) of broadband noise in experiment 1 and for the discrimination of fundamental frequency (F0) of broadband sine-phase harmonic complexes in experiment 2. The reference AMR was 100 Hz as was the reference F0. A child-friendly interface helped listeners to remain attentive to the task. Data were fitted using a maximum-likelihood technique that extracted threshold, slope, and lapse rate. All thresholds were subsequently standardized to a common d' value equal to 0.77. There were relatively large individual differences across listeners: eight had relatively adult-like thresholds in both tasks and nine had higher thresholds. However, these individual differences did not vary systematically with age, over the span of 6-16 yr. Thresholds were correlated across the two tasks and were about nine times finer for F0 discrimination than for AMR discrimination as has been previously observed in adults.

Detection and rate discrimination of amplitude modulation in electrical hearing

Three experiments were designed to examine temporal envelope processing by cochlear implant (CI) listeners. In experiment 1, the hypothesis that listeners' modulation sensitivity would in part determine their ability to discriminate between temporal modulation rates was examined. Temporal modulation transfer functions (TMTFs) obtained in an amplitude modulation detection (AMD) task were compared to threshold functions obtained in an amplitude modulation rate discrimination (AMRD) task. Statistically significant nonlinear correlations were observed between the two measures. In experiment 2, results of loudness-balancing showed small increases in the loudness of modulated over unmodulated stimuli beyond a modulation depth of 16%. Results of experiment 3 indicated small but statistically significant effects of level-roving on the overall gain of the TMTF, but no impact of level-roving on the average shape of the TMTF across subjects. This suggested that level-roving simply increased the task difficulty for most listeners, but did not indicate increased use of intensity cues under more challenging conditions. Data obtained with one subject, however, suggested that the most sensitive listeners may derive some benefit from intensity cues in these tasks. Overall, results indicated that intensity cues did not play an important role in temporal envelope processing by the average CI listener.

Modulation rate discrimination using half-wave rectified and sinusoidally amplitude modulated stimuli in cochlear-implant users

Detection and modulation rate discrimination were measured in cochlear-implant users for pulse-trains that were either sinusoidally amplitude modulated or were modulated with half-wave rectified sinusoids, which in acoustic hearing have been used to simulate the response to low-frequency temporal fine structure. In contrast to comparable results from acoustic hearing, modulation rate discrimination was not statistically different for the two stimulus types. The results suggest that, in contrast to binaural perception, pitch perception in cochlear-implant users does not benefit from using stimuli designed to more closely simulate the cochlear response to low-frequency pure tones.

Factors affecting the use of noise-band vocoders as acoustic models for pitch perception in cochlear implants

Although in a number of experiments noise-band vocoders have been shown to provide acoustic models for speech perception in cochlear implants (CI), the present study assesses in four experiments whether and under what limitations noise-band vocoders can be used as an acoustic model for pitch perception in CI. The first two experiments examine the effect of spectral smearing on simulated electrode discrimination and fundamental frequency (FO) discrimination. The third experiment assesses the effect of spectral mismatch in an FO-discrimination task with two different vocoders. The fourth experiment investigates the effect of amplitude compression on modulation rate discrimination. For each experiment, the results obtained from normal-hearing subjects presented with vocoded stimuli are compared to results obtained directly from CI recipients. The results show that place pitch sensitivity drops with increased spectral smearing and that place pitch cues for multi-channel stimuli can adequately be mimicked when the discriminability of adjacent channels is adjusted by varying the spectral slopes to match that of CI subjects. The results also indicate that temporal pitch sensitivity is limited for noise-band carriers with low center frequencies and that the absence of a compression function in the vocoder might alter the saliency of the temporal pitch cues.

Learning in discrimination of frequency or modulation rate: generalization to fundamental frequency discrimination

Fifteen initially inexperienced subjects were trained for 4 weeks (12 2-h sessions) in frequency discrimination with pure tones around 88, 250, or 1605 Hz, or amplitude modulation rate discrimination of noise bands, using modulation rates around 88 or 250 Hz. Before, in the middle of, and after this training period, pure-tone frequency discrimination thresholds (DLFs), harmonic complex tone fundamental frequency discrimination thresholds (DLF0s), and amplitude modulation rate discrimination thresholds (DLFMs) were measured in several conditions including the trained one. Training in pure-tone frequency discrimination resulted in significantly larger improvements in DLF0s when the test complexes contained resolved harmonics than when they were composed of unresolved harmonics. This result supports the hypothesis that the discrimination of the F0 of resolved harmonics shares common underlying mechanisms with the frequency discrimination of pure tones. Training in rate discrimination did not result in larger DLF0 improvements for unresolved than for resolved harmonics.

Perceptual learning in frequency discrimination and amplitude-modulation rate discrimination, and generalization to fundamental frequency discrimination

Fifteen subjects were trained during twelve 2-h sessions in either frequency discrimination with pure tones, or amplitude-modulation rate discrimination of noise bands. Thresholds for the discrimination of pure-tone frequency, harmonic complex tone fundamental frequency, and amplitude-modulation rate were measured before, during, and after training. Comparison of pre- and post-training thresholds revealed significant improvements in all conditions in both subjects trained in frequency discrimination and subjects trained in modulation rate discrimination. Training in frequency discrimination resulted in larger improvements in fundamental frequency discrimination when the test complexes contained resolved harmonics than when they were composed of unresolved harmonics. Training in modulation rate discrimination did not result in larger fundamental frequency discrimination improvements for unresolved than for resolved harmonics. The implications for models of pitch perception are discussed.

Modulation rate discrimination for unresolved components: temporal cues related to fine structure and envelope.

The present study investigated the hypothesis that the cues for modulation rate discrimination for unresolved spectral components differ as a function of the spectral region occupied by the stimuli. Specifically, it was hypothesized that when components occupy relatively low spectral regions, phase locking both to the fine structure and to the envelope are useful cues. However, as the spectral region occupied by the components increases, phase locking to the fine structure becomes less robust, whereas phase locking to the envelope remains as a potentially strong cue. Observers were asked to detect a decrease in modulation rate for carrier frequencies between 1500 and 6000 Hz. Both amplitude-modulated (AM) and quasifrequency-modulated (QFM) tones were used in order to produce stimuli having strong and weak envelope cues, respectively. Although there were marked individual differences, the results showed an interaction between modulation type and spectral region, with AM and QFM performance being relatively similar at low spectral region, but with QFM showing a steeper reduction in performance as the spectral region of the carrier frequency increased. Overall, the data are consistent with an interpretation that pitch perception for unresolved components depends upon both fine structure and envelope cues, and that the relative importance of these cues depends upon the spectral region occupied by the stimuli.

Modulation rate-discrimination and frequency modulation detection thresholds in the amplitude modulation domain

The present study tested the existence of modulation filters by transposing the concept of excitation pattern to the amplitude modulation (AM) domain. In the first experiment, AM rate-discrimination thresholds (DLFs) for white-noise carriers with AM reference rates of 4, 16, and 64 Hz were measured in five normal-hearing listeners. AM depth was either (i) fixed at 0.2, 0.6, or 1.0, or (ii) varied randomly between 0.2 and 1.0. Randomization of the modulation depth occurred either between trials or between the two intervals of the same trial. In a second experiment, detection thresholds of slow (1 or 4 Hz) frequency modulation (FMDLs) applied to the AM carrier (of identical rates as in the previous experiment) were measured, with and without the presence of a slow sinusoidal modulation applied to the depth of the AM carrier. Overall, the data show that DLFs are less affected than FMDLs by variations in the depth of AM. The results are discussed in light of current models of AM perception.