Human Primary Auditory Cortex Research Articles

Attractor-Like Dynamics Extracted from Human Electrocorticographic Recordings Underlie Computational Principles of Auditory Bistable Perception.

In bistable perception, observers experience alternations between two interpretations of an unchanging stimulus. Neurophysiological studies of bistable perception typically partition neural measurements into stimulus-based epochs and assess neuronal differences between epochs based on subjects' perceptual reports. Computational studies replicate statistical properties of percept durations with modeling principles like competitive attractors or Bayesian inference. However, bridging neuro-behavioral findings with modeling theory requires the analysis of single-trial dynamic data. Here, we propose an algorithm for extracting nonstationary timeseries features from single-trial electrocorticography (ECoG) data. We applied the proposed algorithm to 5-min ECoG recordings from human primary auditory cortex obtained during perceptual alternations in an auditory triplet streaming task (six subjects: four male, two female). We report two ensembles of emergent neuronal features in all trial blocks. One ensemble consists of periodic functions that encode a stereotypical response to the stimulus. The other comprises more transient features and encodes dynamics associated with bistable perception at multiple time scales: minutes (within-trial alternations), seconds (duration of individual percepts), and milliseconds (switches between percepts). Within the second ensemble, we identified a slowly drifting rhythm that correlates with the perceptual states and several oscillators with phase shifts near perceptual switches. Projections of single-trial ECoG data onto these features establish low-dimensional attractor-like geometric structures invariant across subjects and stimulus types. These findings provide supporting neural evidence for computational models with oscillatory-driven attractor-based principles. The feature extraction techniques described here generalize across recording modality and are appropriate when hypothesized low-dimensional dynamics characterize an underlying neural system.SIGNIFICANCE STATEMENT Irrespective of the sensory modality, neurophysiological studies of multistable perception have typically investigated events time-locked to the perceptual switching rather than the time course of the perceptual states per se. Here, we propose an algorithm that extracts neuronal features of bistable auditory perception from largescale single-trial data while remaining agnostic to the subject's perceptual reports. The algorithm captures the dynamics of perception at multiple timescales, minutes (within-trial alternations), seconds (durations of individual percepts), and milliseconds (timing of switches), and distinguishes attributes of neural encoding of the stimulus from those encoding the perceptual states. Finally, our analysis identifies a set of latent variables that exhibit alternating dynamics along a low-dimensional manifold, similar to trajectories in attractor-based models for perceptual bistability.

Quantification of the Intrinsic T1 and T2 of Heschl’s Gyri with MR Fingerprinting

The human primary auditory cortex is located in the Heschl's gyrus (HG). To assess the intrinsic MR property in the gray matter of the HG (GM-HG) with T1 and T2 values using a commercially available MR fingerprinting (MRF) technique. The subjects were 10 healthy volunteers (with 20 HGs; mean age, 31.5 years old; range, 25-53 years old). Coronal T1 and T2 maps were obtained with commercially available MRF using a 3-Tesla MR system. Two radiologists measured the T1 and T2 values of the GM-HG, the GM in the superior temporal gyrus (GM-STG), and the GM in the middle temporal gyrus (GM-MTG) by drawing a ROI on coronal maps. For both radiologists, the mean T1 and T2 values of the GM-HG were significantly lower than those in the GM-STG or GM-MTG (P < 0.01). The interobserver reliability using the intraclass correlation coefficients (ICC) (2,1) showed strong agreement for the measurement of the T1 and T2 values (ICCs =⃥ 0.80 and 0.78 for T1 and T2 values, respectively). The T1 and T2 values on MRF for the GM-HG were lower than those for the GM-STG and GM-MTG, likely reflecting a higher myelin content and iron deposition in the GM-HG. Quantitative measurements using the MRF can clarify cortical properties with high reliability, which may indicate that MRF mapping provides new insights into the structure of the human cortical GM.

Speech Intelligibility Prediction using Spectro-Temporal Modulation Analysis.

Spectro-temporal modulations are believed to mediate the analysis of speech sounds in the human primary auditory cortex. Inspired by humans' robustness in comprehending speech in challenging acoustic environments, we propose an intrusive speech intelligibility prediction (SIP) algorithm, wSTMI, for normal-hearing listeners based on spectro-temporal modulation analysis (STMA) of the clean and degraded speech signals. In the STMA, each of 55 modulation frequency channels contributes an intermediate intelligibility measure. A sparse linear model with parameters optimized using Lasso regression results in combining the intermediate measures of 8 of the most salient channels for SIP. In comparison with a suite of 10 SIP algorithms, wSTMI performs consistently well across 13 datasets, which together cover degradation conditions including modulated noise, noise reduction processing, reverberation, near-end listening enhancement, and speech interruption. We show that the optimized parameters of wSTMI may be interpreted in terms of modulation transfer functions of the human auditory system. Thus, the proposed approach offers evidence affirming previous studies of the perceptual characteristics underlying speech signal intelligibility.

Processing complexity increases in superficial layers of human primary auditory cortex

The layers of the neocortex each have a unique anatomical connectivity and functional role. Their exploration in the human brain, however, has been severely restricted by the limited spatial resolution of non-invasive measurement techniques. Here, we exploit the sensitivity and specificity of ultra-high field fMRI at 7 Tesla to investigate responses to natural sounds at deep, middle, and superficial cortical depths of the human auditory cortex. Specifically, we compare the performance of computational models that represent different hypotheses on sound processing inside and outside the primary auditory cortex (PAC). We observe that while BOLD responses in deep and middle PAC layers are equally well represented by a simple frequency model and a more complex spectrotemporal modulation model, responses in superficial PAC are better represented by the more complex model. This indicates an increase in processing complexity in superficial PAC, which remains present throughout cortical depths in the non-primary auditory cortex. These results suggest that a relevant transformation in sound processing takes place between the thalamo-recipient middle PAC layers and superficial PAC. This transformation may be a first computational step towards sound abstraction and perception, serving to form an increasingly more complex representation of the physical input.

Reduced cortical thickness in Heschl's gyrus as an in vivo marker for human primary auditory cortex

The primary auditory cortex (PAC) is located in the region of Heschl's gyrus (HG), as confirmed by histological, cytoarchitectonical, and neurofunctional studies. Applying cortical thickness (CTH) analysis based on high-resolution magnetic resonance imaging (MRI) and magnetoencephalography (MEG) in 60 primary school children and 60 adults, we investigated the CTH distribution of left and right auditory cortex (AC) and primary auditory source activity at the group and individual level. Both groups showed contoured regions of reduced auditory cortex (redAC) along the mediolateral extension of HG, illustrating large inter-individual variability with respect to shape, localization, and lateralization. In the right hemisphere, redAC localized more within the medial portion of HG, extending typically across HG duplications. In the left hemisphere, redAC was distributed significantly more laterally, reaching toward the anterolateral portion of HG. In both hemispheres, redAC was found to be significantly thinner (mean CTH of 2.34 mm) as compared to surrounding areas (2.99 mm). This effect was more dominant in the right hemisphere rather than in the left one. Moreover, localization of the primary component of auditory evoked activity (P1), as measured by MEG in response to complex harmonic sounds, strictly co-localized with redAC. This structure-function link was found consistently at the group and individual level, suggesting PAC to be represented by areas of reduced cortex in HG. Thus, we propose reduced CTH as an in vivo marker for identifying shape and localization of PAC in the individual brain.

Active Sound Localization Sharpens Spatial Tuning in Human Primary Auditory Cortex.

Spatial hearing sensitivity in humans is dynamic and task-dependent, but the mechanisms in human auditory cortex that enable dynamic sound location encoding remain unclear. Using functional magnetic resonance imaging (fMRI), we assessed how active behavior affects encoding of sound location (azimuth) in primary auditory cortical areas and planum temporale (PT). According to the hierarchical model of auditory processing and cortical functional specialization, PT is implicated in sound location ("where") processing. Yet, our results show that spatial tuning profiles in primary auditory cortical areas (left primary core and right caudo-medial belt) sharpened during a sound localization ("where") task compared with a sound identification ("what") task. In contrast, spatial tuning in PT was sharp but did not vary with task performance. We further applied a population pattern decoder to the measured fMRI activity patterns, which confirmed the task-dependent effects in the left core: sound location estimates from fMRI patterns measured during active sound localization were most accurate. In PT, decoding accuracy was not modulated by task performance. These results indicate that changes of population activity in human primary auditory areas reflect dynamic and task-dependent processing of sound location. As such, our findings suggest that the hierarchical model of auditory processing may need to be revised to include an interaction between primary and functionally specialized areas depending on behavioral requirements.SIGNIFICANCE STATEMENT According to a purely hierarchical view, cortical auditory processing consists of a series of analysis stages from sensory (acoustic) processing in primary auditory cortex to specialized processing in higher-order areas. Posterior-dorsal cortical auditory areas, planum temporale (PT) in humans, are considered to be functionally specialized for spatial processing. However, this model is based mostly on passive listening studies. Our results provide compelling evidence that active behavior (sound localization) sharpens spatial selectivity in primary auditory cortex, whereas spatial tuning in functionally specialized areas (PT) is narrow but task-invariant. These findings suggest that the hierarchical view of cortical functional specialization needs to be extended: our data indicate that active behavior involves feedback projections from higher-order regions to primary auditory cortex.

Frequency-specific attentional modulation in human primary auditory cortex and midbrain

Paying selective attention to an audio frequency selectively enhances activity within primary auditory cortex (PAC) at the tonotopic site (frequency channel) representing that frequency. Animal PAC neurons achieve this ‘frequency-specific attentional spotlight’ by adapting their frequency tuning, yet comparable evidence in humans is scarce. Moreover, whether the spotlight operates in human midbrain is unknown.To address these issues, we studied the spectral tuning of frequency channels in human PAC and inferior colliculus (IC), using 7-T functional magnetic resonance imaging (FMRI) and frequency mapping, while participants focused on different frequency-specific sounds. We found that shifts in frequency-specific attention alter the response gain, but not tuning profile, of PAC frequency channels. The gain modulation was strongest in low-frequency channels and varied near-monotonically across the tonotopic axis, giving rise to the attentional spotlight. We observed less prominent, non-tonotopic spatial patterns of attentional modulation in IC.These results indicate that the frequency-specific attentional spotlight in human PAC as measured with FMRI arises primarily from tonotopic gain modulation, rather than adapted frequency tuning. Moreover, frequency-specific attentional modulation of afferent sound processing in human IC seems to be considerably weaker, suggesting that the spotlight diminishes toward this lower-order processing stage. Our study sheds light on how the human auditory pathway adapts to the different demands of selective hearing.

Differential Rates of Perinatal Maturation of Human Primary and Nonprimary Auditory Cortex.

Primary and nonprimary cerebral cortex mature along different timescales; however, the differences between the rates of maturation of primary and nonprimary cortex are unclear. Cortical maturation can be measured through changes in tissue microstructure detectable by diffusion magnetic resonance imaging (MRI). In this study, diffusion tensor imaging (DTI) was used to characterize the maturation of Heschl’s gyrus (HG), which contains both primary auditory cortex (pAC) and nonprimary auditory cortex (nAC), in 90 preterm infants between 26 and 42 weeks postmenstrual age (PMA). The preterm infants were in different acoustical environments during their hospitalization: 46 in open ward beds and 44 in single rooms. A control group consisted of 15 term-born infants. Diffusion parameters revealed that (1) changes in cortical microstructure that accompany cortical maturation had largely already occurred in pAC by 28 weeks PMA, and (2) rapid changes were taking place in nAC between 26 and 42 weeks PMA. At term equivalent PMA, diffusion parameters for auditory cortex were different between preterm infants and term control infants, reflecting either delayed maturation or injury. No effect of room type was observed. For the preterm group, disturbed maturation of nonprimary (but not primary) auditory cortex was associated with poorer language performance at age two years.

Reconstructing Tone Sequences from Functional Magnetic Resonance Imaging Blood-Oxygen Level Dependent Responses within Human Primary Auditory Cortex.

Here we show that, using functional magnetic resonance imaging (fMRI) blood-oxygen level dependent (BOLD) responses in human primary auditory cortex, it is possible to reconstruct the sequence of tones that a person has been listening to over time. First, we characterized the tonotopic organization of each subject’s auditory cortex by measuring auditory responses to randomized pure tone stimuli and modeling the frequency tuning of each fMRI voxel as a Gaussian in log frequency space. Then, we tested our model by examining its ability to work in reverse. Auditory responses were re-collected in the same subjects, except this time they listened to sequences of frequencies taken from simple songs (e.g., “Somewhere Over the Rainbow”). By finding the frequency that minimized the difference between the model’s prediction of BOLD responses and actual BOLD responses, we were able to reconstruct tone sequences, with mean frequency estimation errors of half an octave or less, and little evidence of systematic biases.

Tonotopic organisation of the auditory cortex in sloping sensorineural hearing loss

Although the tonotopic organisation of the human primary auditory cortex (PAC) has already been studied, the question how its responses are affected in sensorineural hearing loss remains open. Twenty six patients (aged 38.1 ± 9.1 years; 12 men) with symmetrical sloping sensorineural hearing loss (SNHL) and 32 age- and gender-matched controls (NH) participated in an fMRI study using a sparse protocol. The stimuli were binaural 8s complex tones with central frequencies of 400 HzCF, 800 HzCF, 1600 HzCF, 3200 HzCF, or 6400 HzCF, presented at 80 dB(C). In NH responses to all frequency ranges were found in bilateral auditory cortices. The outcomes of a winnermap approach, showing a relative arrangement of active frequency-specific areas, was in line with the existing literature and revealed a V-shape high-frequency gradient surrounding areas that responded to low frequencies in the auditory cortex. In SNHL frequency-specific auditory cortex responses were observed only for sounds from 400 HzCF to 1600 HzCF, due to the severe or profound hearing loss in higher frequency ranges. Using a stringent statistical threshold (p < 0.05; FWE) significant differences between NH and SNHL were only revealed for mid and high-frequency sounds. At a more lenient statistical threshold (p < 0.001, FDRc), however, the size of activation induced by 400 HzCF in PAC was found statistically larger in patients with a prelingual, as compared to a postlingual onset of hearing loss. In addition, this low-frequency range was more extensively represented in the auditory cortex when outcomes obtained in all patients were contrasted with those revealed in normal hearing individuals (although statistically significant only for the secondary auditory cortex). The outcomes of the study suggest preserved patterns of large-scale tonotopic organisation in SNHL which can be further refined following auditory experience, especially when the hearing loss occurs prelingually. SNHL can induce both enlargement and reduction of the extent of responses in the topically organized auditory cortex.

Frequency-Selective Attention in Auditory Scenes Recruits Frequency Representations Throughout Human Superior Temporal Cortex.

A sound of interest may be tracked amid other salient sounds by focusing attention on its characteristic features including its frequency. Functional magnetic resonance imaging findings have indicated that frequency representations in human primary auditory cortex (AC) contribute to this feat. However, attentional modulations were examined at relatively low spatial and spectral resolutions, and frequency-selective contributions outside the primary AC could not be established. To address these issues, we compared blood oxygenation level-dependent (BOLD) responses in the superior temporal cortex of human listeners while they identified single frequencies versus listened selectively for various frequencies within a multifrequency scene. Using best-frequency mapping, we observed that the detailed spatial layout of attention-induced BOLD response enhancements in primary AC follows the tonotopy of stimulus-driven frequency representations-analogous to the "spotlight" of attention enhancing visuospatial representations in retinotopic visual cortex. Moreover, using an algorithm trained to discriminate stimulus-driven frequency representations, we could successfully decode the focus of frequency-selective attention from listeners' BOLD response patterns in nonprimary AC. Our results indicate that the human brain facilitates selective listening to a frequency of interest in a scene by reinforcing the fine-grained activity pattern throughout the entire superior temporal cortex that would be evoked if that frequency was present alone.

Frequency preference and attention effects across cortical depths in the human primary auditory cortex

Columnar arrangements of neurons with similar preference have been suggested as the fundamental processing units of the cerebral cortex. Within these columnar arrangements, feed-forward information enters at middle cortical layers whereas feedback information arrives at superficial and deep layers. This interplay of feed-forward and feedback processing is at the core of perception and behavior. Here we provide in vivo evidence consistent with a columnar organization of the processing of sound frequency in the human auditory cortex. We measure submillimeter functional responses to sound frequency sweeps at high magnetic fields (7 tesla) and show that frequency preference is stable through cortical depth in primary auditory cortex. Furthermore, we demonstrate that-in this highly columnar cortex-task demands sharpen the frequency tuning in superficial cortical layers more than in middle or deep layers. These findings are pivotal to understanding mechanisms of neural information processing and flow during the active perception of sounds.

Decoding speech perception from single cell activity in humans

Deciphering the content of continuous speech is a challenging task performed daily by the human brain. Here, we tested whether activity of single cells in auditory cortex could be used to support such a task. We recorded neural activity from auditory cortex of two neurosurgical patients while presented with a short video segment containing speech. Population spiking activity (~20 cells per patient) allowed detection of word onset and decoding the identity of perceived words with significantly high accuracy levels. Oscillation phase of local field potentials (8–12Hz) also allowed decoding word identity although with lower accuracy levels. Our results provide evidence that the spiking activity of a relatively small population of cells in human primary auditory cortex contains significant information for classification of words in ongoing speech. Given previous evidence for overlapping neural representation during speech perception and production, this may have implications for developing brain–machine interfaces for patients with deficits in speech production.

Population receptive field estimates of human auditory cortex

Here we describe a method for measuring tonotopic maps and estimating bandwidth for voxels in human primary auditory cortex (PAC) using a modification of the population Receptive Field (pRF) model, developed for retinotopic mapping in visual cortex by Dumoulin and Wandell (2008). The pRF method reliably estimates tonotopic maps in the presence of acoustic scanner noise, and has two advantages over phase-encoding techniques. First, the stimulus design is flexible and need not be a frequency progression, thereby reducing biases due to habituation, expectation, and estimation artifacts, as well as reducing the effects of spatio-temporal BOLD nonlinearities. Second, the pRF method can provide estimates of bandwidth as a function of frequency. We find that bandwidth estimates are narrower for voxels within the PAC than in surrounding auditory responsive regions (non-PAC).

Spatial representations of temporal and spectral sound cues in human auditory cortex

Natural and behaviorally relevant sounds are characterized by temporal modulations of their waveforms, which carry important cues for sound segmentation and communication. Still, there is little consensus as to how this temporal information is represented in auditory cortex. Here, by using functional magnetic resonance imaging (fMRI) optimized for studying the auditory system, we report the existence of a topographically ordered spatial representation of temporal sound modulation rates in human auditory cortex. We found a topographically organized sensitivity within auditory cortex to sounds with varying modulation rates, with enhanced responses to lower modulation rates (2 and 4 Hz) on lateral parts of Heschl's gyrus (HG) and faster modulation rates (16 and 32 Hz) on medial HG. The representation of temporal modulation rates was distinct from the representation of sound frequencies (tonotopy) that was orientated roughly orthogonal. Moreover, the combination of probabilistic anatomical maps with a previously proposed functional delineation of auditory fields revealed that the distinct maps of temporal and spectral sound features both prevail within two presumed primary auditory fields hA1 and hR. Our results reveal a topographically ordered representation of temporal sound cues in human primary auditory cortex that is complementary to maps of spectral cues. They thereby enhance our understanding of the functional parcellation and organization of auditory cortical processing.

Tuning In to Sound: Frequency-Selective Attentional Filter in Human Primary Auditory Cortex

Cocktail parties, busy streets, and other noisy environments pose a difficult challenge to the auditory system: how to focus attention on selected sounds while ignoring others? Neurons of primary auditory cortex, many of which are sharply tuned to sound frequency, could help solve this problem by filtering selected sound information based on frequency-content. To investigate whether this occurs, we used high-resolution fMRI at 7 tesla to map the fine-scale frequency-tuning (1.5 mm isotropic resolution) of primary auditory areas A1 and R in six human participants. Then, in a selective attention experiment, participants heard low (250 Hz)- and high (4000 Hz)-frequency streams of tones presented at the same time (dual-stream) and were instructed to focus attention onto one stream versus the other, switching back and forth every 30 s. Attention to low-frequency tones enhanced neural responses within low-frequency-tuned voxels relative to high, and when attention switched the pattern quickly reversed. Thus, like a radio, human primary auditory cortex is able to tune into attended frequency channels and can switch channels on demand.

Altered Cross-Modal Processing in the Primary Auditory Cortex of Congenitally Deaf Adults: A Visual-Somatosensory fMRI Study with a Double-Flash Illusion

The developing brain responds to the environment by using statistical correlations in input to guide functional and structural changes-that is, the brain displays neuroplasticity. Experience shapes brain development throughout life, but neuroplasticity is variable from one brain system to another. How does the early loss of a sensory modality affect this complex process? We examined cross-modal neuroplasticity in anatomically defined subregions of Heschl's gyrus, the site of human primary auditory cortex, in congenitally deaf humans by measuring the fMRI signal change in response to spatially coregistered visual, somatosensory, and bimodal stimuli. In the deaf Heschl's gyrus, signal change was greater for somatosensory and bimodal stimuli than that of hearing participants. Visual responses in Heschl's gyrus, larger in deaf than hearing, were smaller than those elicited by somatosensory stimulation. In contrast to Heschl's gyrus, in the superior-temporal cortex visual signal was comparable to somatosensory signal. In addition, deaf adults perceived bimodal stimuli differently; in contrast to hearing adults, they were susceptible to a double-flash visual illusion induced by two touches to the face. Somatosensory and bimodal signal change in rostrolateral Heschl's gyrus predicted the strength of the visual illusion in the deaf adults in line with the interpretation that the illusion is a functional consequence of the altered cross-modal organization observed in deaf auditory cortex. Our results demonstrate that congenital and profound deafness alters how vision and somatosensation are processed in primary auditory cortex.

Correlates of perceptual awareness in human primary auditory cortex revealed by an informational masking experiment

The presence of an auditory event may remain undetected in crowded environments, even when it is well above the sensory threshold. This effect, commonly known as informational masking, allows for isolating neural activity related to perceptual awareness, by comparing repetitions of the same physical stimulus where the target is either detected or not. Evidence from magnetoencephalography (MEG) suggests that auditory-cortex activity in the latency range 50-250 ms is closely coupled with perceptual awareness. Here, BOLD fMRI and MEG were combined to investigate at which stage in the auditory cortex neural correlates of conscious auditory perception can be observed. Participants were asked to indicate the perception of a regularly repeating target tone, embedded within a random multi-tone masking background. Results revealed widespread activation within the auditory cortex for detected target tones, which was delayed but otherwise similar to the activation of an unmasked control stimulus. The contrast of detected versus undetected targets revealed activity confined to medial Heschl's gyrus, where the primary auditory cortex is located. These results suggest that activity related to conscious perception involves the primary auditory cortex and is not restricted to activity in secondary areas.

Human Primary Auditory Cortex Follows the Shape of Heschl's Gyrus

The primary auditory cortex (PAC) is central to human auditory abilities, yet its location in the brain remains unclear. We measured the two largest tonotopic subfields of PAC (hA1 and hR) using high-resolution functional MRI at 7 T relative to the underlying anatomy of Heschl's gyrus (HG) in 10 individual human subjects. The data reveals a clear anatomical-functional relationship that, for the first time, indicates the location of PAC across the range of common morphological variants of HG (single gyri, partial duplications, and complete duplications). In 20/20 individual hemispheres, two primary mirror-symmetric tonotopic maps were clearly observed with gradients perpendicular to HG. PAC spanned both divisions of HG in cases of partial and complete duplications (11/20 hemispheres), not only the anterior division as commonly assumed. Specifically, the central union of the two primary maps (the hA1-R border) was consistently centered on the full Heschl's structure: on the gyral crown of single HGs and within the sulcal divide of duplicated HGs. The anatomical-functional variants of PAC appear to be part of a continuum, rather than distinct subtypes. These findings significantly revise HG as a marker for human PAC and suggest that tonotopic maps may have shaped HG during human evolution. Tonotopic mappings were based on only 16 min of fMRI data acquisition, so these methods can be used as an initial mapping step in future experiments designed to probe the function of specific auditory fields.

Acoustic experience but not attention modifies neural population phase expressed in human primary auditory cortex

We studied the effect of auditory training on the 40-Hz auditory steady-state response (ASSR) known to localize tonotopically to the region of primary auditory cortex (A1). The stimulus procedure was designed to minimize competitive interactions among frequency representations in A1 and delivered target events at random times in a training window, to increase the likelihood that neuroplastic changes could be detected. Experiment 1 found that repeated exposure to this stimulus advanced the phase of the ASSR (shortened the time delay between the 40-Hz response and stimulus waveforms). The phase advance appeared at the outset of the second of two sessions separated by 24–72 h, did not require active training, and was not accompanied by changes in ASSR amplitude over this time interval. Experiment 2 applied training for 10 sessions to reveal further advances in ASSR phase and also an increase in ASSR amplitude, but the amplitude effect lagged that on phase and did not correlate with perceptual performance while the phase advance did. A control group trained for a single session showed a phase advance but no amplitude enhancement when tested 6 weeks later (retention). In both experiments attention to auditory signals increased ASSR amplitude but had no effect on ASSR phase. Our results reveal a persistent form of neural plasticity expressed in the phase of ASSRs generated from the region of A1, which occurs either in A1 or in subcortical nuclei projecting to this region.