Neural encoding of musical expectations in a non-human primate.

The appreciation of music is a universal trait of humankind.1,2,3 Evidence supporting this notion includes the ubiquity of music across cultures4,5,6,7 and the natural predisposition toward music that humans display early in development.8,9,10 Are we musical animals because of species-specific predispositions? This question cannot be answered by relying on cross-cultural or developmental studies alone, as these cannot rule out enculturation.11 Instead, it calls for cross-species experiments testing whether homologous neural mechanisms underlying music perception are present in non-human primates. We present music to two rhesus monkeys, reared without musical exposure, while recording electroencephalography (EEG) and pupillometry. Monkeys exhibit higher engagement and neural encoding of expectations based on the previously seeded musical context when passively listening to real music as opposed to shuffled controls. We then compare human and monkey neural responses to the same stimuli and find a species-dependent contribution of two fundamental musical features-pitch and timing12-in generating expectations: while timing- and pitch-based expectations13 are similarly weighted in humans, monkeys rely on timing rather than pitch. Together, these results shed light on the phylogeny of music perception. They highlight monkeys' capacity for processing temporal structures beyond plain acoustic processing, and they identify a species-dependent contribution of time- and pitch-related features to the neural encoding of musical expectations.

The influence of musical parameters and subjective musical ratings on perceptions of culture.

Recent research suggests that music can affect evaluations of other groups and cultures. However, little is known about the objective and subjective musical parameters that influence these evaluations. We aimed to fill this gap through two studies. Study 1 collected responses from 52 American participants who listened to 30 folk-song melodies from different parts of the world. Linear mixed-effects models tested the influence of objective and subjective musical parameters of these melodies on evaluations of the cultures from which they originated. Musical parameters consistently predicted cultural evaluations. The most prominent musical parameter was musical velocity, a measure of number of pitch onsets, predicting more cultural warmth, competence and evolvedness and less cultural threat. Next, with a sample of 212 American participants, Study 2 used a within-subjects experiment to alter the tempo and dissonance for a subset of six melody excerpts from Study 1, testing for causal effects. Linear mixed-effects models revealed that both dissonance and slow tempo predicted more negative cultural evaluations. Together, both studies demonstrate how musical parameters can influence cultural perceptions. Avenues for future research are discussed.

Neurophysiological time course of timbre-induced music-like perception.

Traditionally, pitch variation in a sound stream has been integral to music identity. We attempt to expand music's definition, by demonstrating that the neural code for musicality is independent of pitch encoding. That is, pitchless sound streams can still induce music-like perception and a neurophysiological hierarchy similar to pitched melodies. Previous work reported that neural processing of sounds with no-pitch, fixed-pitch, and irregular-pitch (melodic) patterns, exhibits a right-lateralized hierarchical shift, with pitchless sounds favorably processed in Heschl's gyrus (HG), ascending laterally to nonprimary auditory areas for fixed-pitch and even more laterally for melodic patterns. The objective of this EEG study was to assess whether sound encoding maintains a similar hierarchical profile when musical perception is driven by timbre irregularities in the absence of pitch changes. Individuals listened to repetitions of three musical and three nonmusical sound-streams. The nonmusical streams were comprised of seven 200-ms segments of white, pink, or brown noise, separated by silent gaps. Musical streams were created similarly, but with all three noise types combined in a unique order within each stream to induce timbre variations and music-like perception. Subjects classified the sound streams as musical or nonmusical. Musical processing exhibited right dominant α power enhancement, followed by a lateralized increase in θ phase-locking and spectral power. The θ phase-locking was stronger in musicians than in nonmusicians. The lateralization of activity suggests higher-level auditory processing. Our findings validate the existence of a hierarchical shift, traditionally observed with pitched-melodic perception, underscoring that musicality can be achieved with timbre irregularities alone.NEW & NOTEWORTHY EEG induced by streams of pitchless noise segments varying in timbre were classified as music-like and exhibited a right-lateralized hierarchy in processing similar to pitched melodic processing. This study provides evidence that the neural-code of musicality is independent of pitch encoding. The results have implications for understanding music processing in individuals with degraded pitch perception, such as in cochlear-implant listeners, as well as the role of nonpitched sounds in the induction of music-like perceptual states.

Articulation posture influences pitch during singing imagery.

Facial muscle activity contributes to singing and to articulation: in articulation, mouth shape can alter vowel identity; and in singing, facial movement correlates with pitch changes. Here, we examine whether mouth posture causally influences pitch during singing imagery. Based on perception-action theories and embodied cognition theories, we predict that mouth posture influences pitch judgments even when no overt utterances are produced. In two experiments (total N = 160), mouth posture was manipulated to resemble the articulation of either /i/ (as in English meet; retracted lips) or /o/ (as in French rose; protruded lips). Holding this mouth posture, participants were instructed to mentally "sing" given songs (which were all positive in valence) while listening with their inner ear and, afterwards, to assess the pitch of their mental chant. As predicted, compared to the o-posture, the i-posture led to higher pitch in mental singing. Thus, bodily states can shape experiential qualities, such as pitch, during imagery. This extends embodied music cognition and demonstrates a new link between language and music.

Distribution of multiunit pitch responses recorded intracranially from human auditory cortex.

The perception of pitch is a fundamental percept, which is mediated by the auditory system, requiring the abstraction of stimulus properties related to the spectro-temporal structure of sound. Despite its importance, there is still debate as to the precise areas responsible for its encoding, which may be due to species differences or differences in the recording measures and choices of stimuli used in previous studies. Moreover, it was unknown whether the human brain contains pitch neurons and how distributed such neurons might be. Here, we present the first study to measure multiunit neural activity in response to pitch stimuli in the auditory cortex of intracranially implanted humans. The stimulus sets were regular-interval noise with a pitch strength that is related to the temporal regularity and a pitch value determined by the repetition rate and harmonic complexes. Specifically, we demonstrate reliable responses to these different pitch-inducing paradigms that are distributed throughout Heschl's gyrus, rather than being localized to a particular region, and this finding was evident regardless of the stimulus presented. These data provide a bridge across animal and human studies and aid our understanding of the processing of a critical percept associated with acoustic stimuli.

Absolute pitch: neurophysiological evidence for early brain activity in prefrontal cortex.

Absolute pitch (AP) is the ability to rapidly label pitch without an external reference. The speed of AP labeling may be related to faster sensory processing. We compared time needed for auditory processing in AP musicians, non-AP musicians, and nonmusicians (NM) using high-density electroencephalographic recording. Participants responded to pure tones and sung voice. Stimuli evoked a negative deflection peaking at ~100 ms (N1) post-stimulus onset, followed by a positive deflection peaking at ~200 ms (P2). N1 latency was shortest in AP, intermediate in non-AP musicians, and longest in NM. Source analyses showed decreased auditory cortex and increased frontal cortex contributions to N1 for complex tones compared with pure tones. Compared with NM, AP musicians had weaker source currents in left auditory cortex but stronger currents in left inferior frontal gyrus (IFG) during N1, and stronger currents in left IFG during P2. Compared with non-AP musicians, AP musicians exhibited stronger source currents in right insula and left IFG during N1, and stronger currents in left IFG during P2. Non-AP musicians had stronger N1 currents in right auditory cortex than nonmusicians. Currents in left IFG and left auditory cortex were correlated to response times exclusively in AP. Findings suggest a left frontotemporal network supports rapid pitch labeling in AP.

The left inferior frontal gyrus is causally linked to vocal feedback control: evidence from high-definition transcranial alternating current stimulation.

Current models of speech motor control propose a role for the left inferior frontal gyrus (IFG) in feedforward control of speech production. There is evidence, however, that has implicated the functional relevance of the left IFG for the neuromotor processing of vocal feedback errors. The present event-related potential (ERP) study examined whether the left IFG is causally linked to auditory feedback control of vocal production with high-definition transcranial alternating current stimulation (HD-tACS). After receiving active or sham HD-tACS over the left IFG at 6 or 70 Hz, 20 healthy adults vocalized the vowel sounds while hearing their voice unexpectedly pitch-shifted by ±200 cents. The results showed that 6 or 70 Hz HD-tACS over the left IFG led to larger magnitudes and longer latencies of vocal compensations for pitch perturbations paralleled by larger ERP P2 responses than sham HD-tACS. Moreover, there was a lack of frequency specificity that showed no significant differences between 6 and 70 Hz HD-tACS. These findings provide first causal evidence linking the left IFG to vocal pitch regulation, suggesting that the left IFG is an important part of the feedback control network that mediates vocal compensations for auditory feedback errors.

Neural correlates of concurrent sound perception: A review and guidelines for future research.

The perception of concurrent sound sources depends on processes (i.e., auditory scene analysis) that fuse and segregate acoustic features according to harmonic relations, temporal coherence, and binaural cues (encompass dichotic pitch, location difference, simulated echo). The object-related negativity (ORN) and P400 are electrophysiological indices of concurrent sound perception. Here, we review the different paradigms used to study concurrent sound perception and the brain responses obtained from these paradigms. Recommendations regarding the design and recording parameters of the ORN and P400 are made, and their clinical applications in assessing central auditory processing ability in different populations are discussed.

Task-modulated Sensitivity to Vocal Pitch in the Dorsal Premotor Cortex during Multitalker Speech Recognition.

It has long been known that listening to speech activates inferior frontal (pre-)motor regions in addition to a more dorsal premotor site (dPM). Recent work shows that dPM, located adjacent to laryngeal motor cortex, responds to low-level acoustic speech cues including vocal pitch, and the speech envelope, in addition to higher-level cues such as phoneme categories. An emerging hypothesis is that dPM is part of a general auditory-guided laryngeal control circuit that plays a role in producing speech and other voluntary auditory-vocal behaviors. We recently reported a study in which dPM responded to vocal pitch during a degraded speech recognition task, but only when speech was rated as unintelligible; dPM was more robustly modulated by the categorical difference between intelligible and unintelligible speech. Contrary to the general auditory-vocal hypothesis, this suggests intelligible speech is the primary driver of dPM. However, the same pattern of results was observed in pitch-sensitive auditory cortex. Crucially, vocal pitch was not relevant to the intelligibility judgment task, which may have facilitated processing of phonetic information at the expense of vocal pitch cues. The present fMRI study (n = 25) tests the hypothesis that, for a multitalker task that emphasizes pitch for talker segregation, left dPM and pitch-sensitive auditory regions will respond to vocal pitch regardless of overall speech intelligibility. This would suggest that pitch processing is indeed a primary concern of this circuit, apparent during perception only when the task demands it. Spectrotemporal modulation distortion was used to independently modulate vocal pitch and phonetic content in two-talker (male/female) utterances across two conditions (Competing, Unison), only one of which required pitch-based segregation (Competing). A Bayesian hierarchical drift-diffusion model was used to predict speech recognition performance from patterns of spectrotemporal distortion imposed on each trial. The model's drift rate parameter, a d'-like measure of performance, was strongly associated with vocal pitch for Competing but not Unison. Using a second Bayesian hierarchical model, we identified regions where behaviorally relevant acoustic features were related to fMRI activation in dPM. We regressed the hierarchical drift-diffusion model's posterior predictions of trial-wise drift rate, reflecting the relative presence or absence of behaviorally relevant acoustic features from trial to trial, against trial-wise activation amplitude. A significant positive association with overall drift rate, reflecting vocal pitch and phonetic cues related to overall intelligibility, was observed in left dPM and bilateral auditory cortex in both conditions. A significant positive association with "pitch-restricted" drift rate, reflecting only the relative presence or absence of behaviorally relevant pitch cues, regardless of the presence or absence of phonetic content (intelligibility), was observed in left dPM, but only in the Competing condition. Interestingly, the same effect was observed in bilateral auditory cortex but in both conditions. A post hoc mediation analysis ruled out the possibility that decision load was responsible for the observed pitch effects. These findings suggest that processing of vocal pitch is a primary concern of the auditory-cortex-dPM circuit, although during perception core pitch, processing is carried out by auditory cortex with a potential modulatory influence from dPM.

Enhanced mismatch negativity in harmonic compared with inharmonic sounds.

Many natural sounds have frequency spectra composed of integer multiples of a fundamental frequency. This property, known as harmonicity, plays an important role in auditory information processing. However, the extent to which harmonicity influences the processing of sound features beyond pitch is still unclear. This is interesting because harmonic sounds have lower information entropy than inharmonic sounds. According to predictive processing accounts of perception, this property could produce more salient neural responses due to the brain's weighting of sensory signals according to their uncertainty. In the present study, we used electroencephalography to investigate brain responses to harmonic and inharmonic sounds commonly occurring in music: Piano tones and hi-hat cymbal sounds. In a multifeature oddball paradigm, we measured mismatch negativity (MMN) and P3a responses to timbre, intensity, and location deviants in listeners with and without congenital amusia-an impairment of pitch processing. As hypothesized, we observed larger amplitudes and earlier latencies (for both MMN and P3a) in harmonic compared with inharmonic sounds. These harmonicity effects were modulated by sound feature. Moreover, the difference in P3a latency between harmonic and inharmonic sounds was larger for controls than amusics. We propose an explanation of these results based on predictive coding and discuss the relationship between harmonicity, information entropy, and precision weighting of prediction errors.

Tonal structures benefit short-term memory for real music: Evidence from non-musicians and individuals with congenital amusia.

Congenital amusia is a neurodevelopmental disorder of music processing, which includes impaired pitch memory, associated to abnormalities in the right fronto-temporal network. Previous research has shown that tonal structures (as defined by the Western musical system) improve short-term memory performance for short tone sequences (in comparison to atonal versions) in non-musician listeners, but the tonal structures only benefited response times in amusic individuals. We here tested the potential benefit of tonal structures for short-term memory with more complex musical material. Congenital amusics and their matched non-musician controls were required to indicate whether two excerpts were the same or different. Results confirmed impaired performance of amusic individuals in this short-term memory task. However, most importantly, both groups of participants showed better memory performance for tonal material than for atonal material. These results revealed that even amusics' impaired short-term memory for pitch shows classical characteristics of short-term memory, that is the mnemonic benefit of structure in the to-be-memorized material. The findings show that amusic individuals have acquired some implicit knowledge of regularities of their culture, allowing for implicit processing of tonal structures, which benefits to memory even for complex material.

Temporal Pitch Sensitivity in an Animal Model: Psychophysics and Scalp Recordings : Temporal Pitch Sensitivity in Cat.

Cochlear implant (CI) users show limited sensitivity to the temporal pitch conveyed by electric stimulation, contributing to impaired perception of music and of speech in noise. Neurophysiological studies in cats suggest that this limitation is due, in part, to poor transmission of the temporal fine structure (TFS) by the brainstem pathways that are activated by electrical cochlear stimulation. It remains unknown, however, how that neural limit might influence perception in the same animal model. For that reason, we developed non-invasive psychophysical and electrophysiological measures of temporal (i.e., non-spectral) pitch processing in the cat. Normal-hearing (NH) cats were presented with acoustic pulse trains consisting of band-limited harmonic complexes that simulated CI stimulation of the basal cochlea while removing cochlear place-of-excitation cues. In the psychophysical procedure, trained cats detected changes from a base pulse rate to a higher pulse rate. In the scalp-recording procedure, the cortical-evoked acoustic change complex (ACC) and brainstem-generated frequency following response (FFR) were recorded simultaneously in sedated cats for pulse trains that alternated between the base and higher rates. The range of perceptual sensitivity to temporal pitch broadly resembled that of humans but was shifted to somewhat higher rates. The ACC largely paralleled these perceptual patterns, validating its use as an objective measure of temporal pitch sensitivity. The phase-locked FFR, in contrast, showed strong brainstem encoding for all tested pulse rates. These measures demonstrate the cat's perceptual sensitivity to pitch in the absence of cochlear-place cues and may be valuable for evaluating neural mechanisms of temporal pitch perception in the feline animal model of stimulation by a CI or novel auditory prostheses.

Principal component decomposition of acoustic and neural representations of time-varying pitch reveals adaptive efficient coding of speech covariation patterns.

Understanding the effects of statistical regularities on speech processing is a central issue in auditory neuroscience. To investigate the effects of distributional covariance on the neural processing of speech features, we introduce and validate a novel approach: decomposition of time-varying signals into patterns of covariation extracted with Principal Component Analysis. We used this decomposition to assay the sensory representation of pitch covariation patterns in native Chinese listeners and non-native learners of Mandarin Chinese tones. Sensory representations were examined using the frequency-following response, a far-field potential that reflects phase-locked activity from neural ensembles along the auditory pathway. We found a more efficient representation of the covariation patterns that accounted for more redundancy in the form of distributional covariance. Notably, long-term language and short-term training experiences enhanced the sensory representation of these covariation patterns.

Cortical activity evoked by voice pitch changes: A combined fNIRS and EEG study.

Functional near-infrared spectroscopy (fNIRS) and electroencephalography (EEG) data were simultaneously obtained from normal-hearing listeners presented with continuous natural vowel sequences to study the interrelation of the haemodynamic and electrophysiological cortical responses evoked by voice pitch changes. fNIRS topographies and distributed ERP source reconstructions both indicated additional activity in the right superior temporal cortex if the prosodic contours varied between successive vowels, rather than being the same throughout the sequences. The source-level ERPs furthermore revealed two temporally and spatially separable adaptation processes in superior temporal cortex: Firstly, the early P1 component was bilaterally attenuated when vowels with the same prosodic contours were presented repeatedly, reflecting sensory adaptation. Secondly, the later P2 and sustained potential components were smaller in the right hemisphere during sequences without prosodic changes, which is taken to represent an attention-based adaptation effect. The present results demonstrate the convergence of both methods and demonstrate which ERPs underlie the right-lateralised activity in superior temporal cortex evoked in response to pitch changes that has been observed in many neuroimaging studies.

Advantages of Pulse Rate Compared to Modulation Frequency for Temporal Pitch Perception in Cochlear Implant Users.

Most cochlear implants encode the fundamental frequency of periodic sounds by amplitude modulation of constant-rate pulsatile stimulation. Pitch perception provided by such stimulation strategies is markedly poor. Two experiments are reported here that consider potential advantages of pulse rate compared to modulation frequency for providing stimulation timing cues for pitch. The first experiment examines beat frequency distortion that occurs when modulating constant-rate pulsatile stimulation. This distortion has been reported on previously, but the results presented here indicate that distortion occurs for higher stimulation rates than previously reported. The second experiment examines pitch resolution as provided by pulse rate compared to modulation frequency. The results indicate that pitch discrimination is better with pulse rate than with modulation frequency. The advantage was large for rates near what has been suggested as the upper limit of temporal pitch perception conveyed by cochlear implants. The results are relevant to sound processing design for cochlear implants particularly for algorithms that encode fundamental frequency into deep envelope modulations or into precisely timed pulsatile stimulation.

Distinct Representations of Tonotopy and Pitch in Human Auditory Cortex.

Frequency-to-place mapping, or tonotopy, is a fundamental organizing principle throughout the auditory system, from the earliest stages of auditory processing in the cochlea to subcortical and cortical regions. Although cortical maps are referred to as tonotopic, it is unclear whether they simply reflect a mapping of physical frequency inherited from the cochlea, a computation of pitch based on the fundamental frequency, or a mixture of these two features. We used high-resolution functional magnetic resonance imaging (fMRI) to measure BOLD responses as male and female human participants listened to pure tones that varied in frequency or complex tones that varied in either spectral content (brightness) or fundamental frequency (pitch). Our results reveal evidence for pitch tuning in bilateral regions that partially overlap with the traditional tonotopic maps of spectral content. In general, primary regions within Heschl's gyri (HGs) exhibited more tuning to spectral content, whereas areas surrounding HGs exhibited more tuning to pitch.SIGNIFICANCE STATEMENT Tonotopy, an orderly mapping of frequency, is observed throughout the auditory system. However, it is not known whether the tonotopy observed in the cortex simply reflects the frequency spectrum (as in the ear) or instead represents the higher-level feature of fundamental frequency, or pitch. Using carefully controlled stimuli and high-resolution functional magnetic resonance imaging (fMRI), we separated these features to study their cortical representations. Our results suggest that tonotopy in primary cortical regions is driven predominantly by frequency, but also reveal evidence for tuning to pitch in regions that partially overlap with the tonotopic gradients but extend into nonprimary cortical areas. In addition to resolving ambiguities surrounding cortical tonotopy, our findings provide evidence that selectivity for pitch is distributed bilaterally throughout auditory cortex.

Longitudinal auditory pathophysiology following mild blast-induced trauma.

Blast-induced hearing difficulties affect thousands of veterans and civilians. The long-term impact of even a mild blast exposure on the central auditory system is hypothesized to contribute to lasting behavioral complaints associated with mild blast traumatic brain injury (bTBI). Although recovery from mild blast has been studied separately over brief or long time windows, few, if any, studies have investigated recovery longitudinally over short-term and longer-term (months) time windows. Specifically, many peripheral measures of auditory function either recover or exhibit subclinical deficits, masking deficits in processing complex, real-world stimuli that may recover differently. Thus, examining the acute time course and pattern of neurophysiological impairment using appropriate stimuli is critical to better understanding and intervening in bTBI-induced auditory system impairments. Here, we compared auditory brainstem response, middle-latency auditory-evoked potentials, and envelope following responses. Stimuli were clicks, tone pips, amplitude-modulated tones in quiet and in noise, and speech-like stimuli (iterated rippled noise pitch contours) in adult male rats subjected to mild blast and sham exposure over the course of 2 mo. We found that blast animals demonstrated drastic threshold increases and auditory transmission deficits immediately after blast exposure, followed by substantial recovery during the window of 7-14 days postblast, although with some deficits remaining even after 2 mo. Challenging conditions and speech-like stimuli can better elucidate mild bTBI-induced auditory deficit during this period. Our results suggest multiphasic recovery and therefore potentially different time windows for treatment, and deficits can be best observed using a small battery of sound stimuli.NEW & NOTEWORTHY Few studies on blast-induced hearing deficits go beyond simple sounds and sparsely track postexposure. Therefore, the recovery arc for potential therapies and real-world listening is poorly understood. Evidence suggested multiple recovery phases over 2 mo postexposure. Hearing thresholds largely recovered within 14 days and partially explained recovery. However, midlatency responses, responses to amplitude modulation in noise, and speech-like pitch sweeps exhibited extended changes, implying persistent central auditory deficits and the importance of subclinical threshold shifts.

Learning nonnative speech sounds changes local encoding in the adult human cortex.

Adults can learn to identify nonnative speech sounds with training, albeit with substantial variability in learning behavior. Increases in behavioral accuracy are associated with increased separability for sound representations in cortical speech areas. However, it remains unclear whether individual auditory neural populations all show the same types of changes with learning, or whether there are heterogeneous encoding patterns. Here, we used high-resolution direct neural recordings to examine local population response patterns, while native English listeners learned to recognize unfamiliar vocal pitch patterns in Mandarin Chinese tones. We found a distributed set of neural populations in bilateral superior temporal gyrus and ventrolateral frontal cortex, where the encoding of Mandarin tones changed throughout training as a function of trial-by-trial accuracy ("learning effect"), including both increases and decreases in the separability of tones. These populations were distinct from populations that showed changes as a function of exposure to the stimuli regardless of trial-by-trial accuracy. These learning effects were driven in part by more variable neural responses to repeated presentations of acoustically identical stimuli. Finally, learning effects could be predicted from speech-evoked activity even before training, suggesting that intrinsic properties of these populations make them amenable to behavior-related changes. Together, these results demonstrate that nonnative speech sound learning involves a wide array of changes in neural representations across a distributed set of brain regions.

Switching perception of musical meters by listening to different acoustic cues of biphasic sound stimulus.

Meter is one of the core features of music perception. It is the cognitive grouping of regular sound sequences, typically for every 2, 3, or 4 beats. Previous studies have suggested that one can not only passively perceive the meter from acoustic cues such as loudness, pitch, and duration of sound elements, but also actively perceive it by paying attention to isochronous sound events without any acoustic cues. Studying the interaction of top-down and bottom-up processing in meter perception leads to understanding the cognitive system's ability to perceive the entire structure of music. The present study aimed to demonstrate that meter perception requires the top-down process (which maintains and switches attention between cues) as well as the bottom-up process for discriminating acoustic cues. We created a "biphasic" sound stimulus, which consists of successive tone sequences designed to provide cues for both the triple and quadruple meters in different sound attributes, frequency, and duration. Participants were asked to focus on either frequency or duration of the stimulus, and to answer how they perceived meters on a five-point scale (ranged from "strongly triple" to "strongly quadruple"). As a result, we found that participants perceived different meters by switching their attention to specific cues. This result adds evidence to the idea that meter perception involves the interaction between top-down and bottom-up processes.

Midbrain-Level Neural Correlates of Behavioral Tone-in-Noise Detection: Dependence on Energy and Envelope Cues.

Hearing in noise is a problem often assumed to depend on encoding of energy level by channels tuned to target frequencies, but few studies have tested this hypothesis. The present study examined neural correlates of behavioral tone-in-noise (TIN) detection in budgerigars (Melopsittacus undulatus, either sex), a parakeet species with human-like behavioral sensitivity to many simple and complex sounds. Behavioral sensitivity to tones in band-limited noise was assessed using operant-conditioning procedures. Neural recordings were made in awake animals from midbrain-level neurons in the inferior colliculus, the first processing stage of the ascending auditory pathway with pronounced rate-based encoding of stimulus amplitude modulation. Budgerigar TIN detection thresholds were similar to human thresholds across the full range of frequencies (0.5-4 kHz) and noise levels (45-85 dB SPL) tested. Also as in humans, thresholds were minimally affected by a challenging roving-level condition with random variation in background-noise level. Many midbrain neurons showed a decreasing response rate as TIN signal-to-noise ratio (SNR) was increased by elevating the tone level, a pattern attributable to amplitude-modulation tuning in these cells and the fact that higher SNR tone-plus-noise stimuli have flatter amplitude envelopes. TIN thresholds of individual neurons were as sensitive as behavioral thresholds under most conditions, perhaps surprisingly even when the unit's characteristic frequency was tuned an octave or more away from the test frequency. A model that combined responses of two cell types enhanced TIN sensitivity in the roving-level condition. These results highlight the importance of midbrain-level envelope encoding and off-frequency neural channels for hearing in noise.SIGNIFICANCE STATEMENT Detection of target sounds in noise is often assumed to depend on energy-level encoding by neural processing channels tuned to the target frequency. In contrast, we found that tone-in-noise sensitivity in budgerigars was often greatest in midbrain neurons not tuned to the test frequency, underscoring the potential importance of off-frequency channels for perception. Furthermore, the results highlight the importance of envelope processing for hearing in noise, especially under challenging conditions with random variation in background noise level over time.